SIDA, contributions to botany.

CONTENTS

The genus Cybianthus (Myrsinaceae) in Ecuador and Peru
Jobn fj. Pipoly HT
1

New flowering plants from meee New Ireland, Papua New Guinea
W. Takeuchi and John J. Pipoly IL
161

Three new species of Crataegus (Rosaceae) from western North America:
C. okennonii, C. okanaganensis and C. phippsii

J.B. Phipps and R.J. O’Kennon

169

Taxonomic clarification of Atriplex nuttallii (Chenopodiaceae) and its near relatives
Howard C. Stutz and Stewart C. Sanderson
193

Flavonoids in Strophostyles species and the related genus Dolichopsis
(Phaseolinae, Fabaceae): Distribution and phylogenetic significance
Juan Pablo Pelotto and Maria A. Del Pero Martinez

213

Ruellia jimulcensis (Acanthaceae), a new species from the Chihuahuan
Desert area, Mexico

José A. Villarreal Q.
223
Una nueva especie de Agave, subgénero Littaea (Agavaceae) de Guerrero y
Oaxaca; México
Abisai Garcia-Mendoza and Esteban Martinez Salas
227
A new species of Mandevilla (Apocynaceae) from Jalisco, Mexico
Justin Williams

231
A new combination in Mexican Mandevilla (Apocynaceae)
Justin Williams
237
mayaénsis (Poaceae: Andropogoneae): A new species from Belize
ra K. Tae and Stanley D. Jones

Taxonomy of the Sporobolus floridanus complex (Poaceae: Sporobolinae)
Alan S. Weakley and Paul M. Peterson
247

Caryopsis morphology of Leptochloa sensu lato| (Poaceae, Chloridoideae)
Neil Snow

CONTRIBUTIONS
TO BOTANY

Xanthostemon fruticosus (Myrtacea€), a mew species from the Philippines
Peter G. Wilson and Leonardo L. Co
283

VOLUME 18
NUMBER 1
OCTOBER, 1998

(continued on back cover)

CONTRIBUTIONS TO BOTANY

FOUNDED BY
LLOYD H. SHINNERS
1962

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Northern Kentucky University Universidad de Leon
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© SIDA, CONTRIBUTIONS TO BOTANY, Volume 18, Number 1, pages 1-376.
opyright 1998
Botanical Research Institute of Texas, Inc.
Printed in the United States of America
ISSN 0036-1488

THE GENUS CYBIANTHUS (MYRSINACEAE)
IN ECUADOR AND PERU

JOHN J. PIPOLY III o

Botanical Research Institute of Texas
9 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
Jpipoly @ brit.org; clusia@latinmail.com

ABSTRACT

The genus Cybianthus was revised to provide taxonomic treatments of the Myrsinaceae
for Flora of Peru, Catalogue of the Vascular Plants of Ecuador, and Flora of Ecuador. Eight of
Cybianthus’ ten subgenera are represented, and an updated description of the genus, keys
to its subgenera and emended descriptions for each are provided. Detailed eon of
the only anatomy and ecology of the genus are presented. Cybianthus subgenus Iteoides
is relegated to synonymy under subgenus Mécroconomorpha. Within each subgenus, Bale
full eee synonymy, distribution, ecology and conservation statuses, local names
and uses are given for each species. In addition, specimens are cited for each species, in-
am reid) ones to show SeeeeS in pinekphologica! vatieton: . Thirteen species

and one, C. is neotypif fied

One new combination, Cybianthus guyanensis anne pseudoicacoreus, : sees and Cybianthus
poeppigii 1 is transferred from subgenus C) ee to subgenus Weige/tia. The following 15

new species are described, illustrated and their phylogenetic relationships are discussed:
Be anthuriophyllus, C. cenepensis, C. Pa i C. croatii, C. flavovirens, C. fosteri
C. grandezit, C. elie C. huampamiensis, C. incognitus, C. gensonti, C. nestorii, C. peD ope fois
C. timanae, C. vasque

RESUMEN

Al preparar tratamientos taxondémicos sobre la familia Myrsinaceae para los proyectos
Flora del Perti, Catdlago de las Plantas Vasculares ae Ecuador, a tee a Ecuador, se llevé a
cabo una revision del género Cybianthus. Se la os diez subgéneros,
y se provéen tanto una descripcién actualizada para el género como para cada subgénero
también, Se presenta SNe oes Saiaeunte tratando la once anatomia y Eee! a

del género = relega C la sinonimiab
y se transfiere Cybiantbus poeppigit del shee ee al subgéneto petit Par
cada subgénero, se f | sinonimia, distribucién geograf

oad

y esta do actual d

nombres locales sett usos. También se cita colecciones revisadas
para cada especie, incl d llas fi de la regi gion delimitada estran variacion
morfolégica ae Se relegan 13 especies a la sinonimia, se eae 9 binomiales
y uno, Conomorpha rigida, se neotipifica. Se publica la nueva combinacén, Cybianthus guyanensis
subsp. pseadoicacoreus. Se describen, se ilustran y se discuten el parentezco para 15 especies
nuevas, listadas a continuacién: ee anthuriophyllus, C. cenepensis, C. comperuvianus,
C. croatii, C. flavovirens, C. i, C. granulosus, C. ie C. incognitus,
C. jensonii, C. nestorii, C. hen ee C timanae, C. vasque

Sipa 18(1): 1-160. 1998

2 Stipa 18(1)
INTRODUCTION

The genus Cybianthus Martius was cladistically defined by the presence
of unique glandular granules at the junction of the corolla tube and lobes
by Pipoly (1987). In addition, the unique combination of lateral (axillary)
inflorescences, gamosepalous and gamopetalous flowers, and stamens con-
nate oy their filaments at least one-fourth their length, and adnate to the
corolla tube at least one-third its length (Pipoly 1987, 1992a) allows for
eee recognition from herbarium specimens. With this contribution,

‘ybianthus now contains 167 species in ten subgenera. The genus includes

species formerly included in the genera Comomyrsine Hook. f., Conomorpha
A. DC., Correlliana D'Arcy, Cybianthopsis Lundell, eis Benth.,
Microconomorpha Lundell, and Wergeltia A. DC. (Pipoly 1987, 1992a). In Ecuador
and Peru, 43 species in 8 subgenera are known. Among the species, none
are endemic to Ecuador, while 11 are endemic to Peru.

The broad generic concept for Cybianthus was first proposed by Agostini
(1970) as a precursor to his dissertation (1972); that was followed by the
formal transfers and a key to the subgenera (Agostini 1980). Subsequently,
while revising subgenus Grammadenia, Pipoly (1987) determined that the
paleotropical genus Embel7a Burman f. was the sister group to the entire
genus and that it was most parsimonius to include Grammadenia within
Cybianthus. Preparation of taxonomic treatments for the Myrsinaceae in Flora
of Peru, Catalogue of the Vascular Plants of Ecuador and Flora of Ecuador re-
vealed that many taxa were new, and much confusion had arisen among the
concepts prevalent for taxa already described. Among adjacent countries
with significant Amazonian regions, Ecuador and Peru share more species
of Cybianthus in common than do any other pair of countries. While it would
have been desirable to include Colombia to provide a treatment for the entire
northwestern South America, the number of additional species endemic to
Colombia, or known only from Colombia and Venezuela, would have in-
creased the size of the treatment by fifty percent. Given that it will be some
time until a revision of the entire genus for F/ora Neotropica is complete, the
present treatment is intended to serve as a precursor to that monograp
and to make the names available for the other aforementioned projects. A
revised description of the genus Cybianthus follows, including keys and emended
descriptions for the eight subgenera occurring in Ecuador and Peru, along
with keys to the taxa and full descriptions for each. For each of the fifteen
new species described, illustrations are provided, while for all species, full
descriptions and synonymy, discussions of distribution, ecology and con-
servation status, etymology, and when known, local names and uses are elucidated.

a

MORPHOLOGY

Morphological terms in this treatment follow Lindley (1848) and Pipoly

Pirpoty, Cybianthus in Ecuador and Peru 3)

(1987, 1992a) for the inflorescence, rachis pedicels and floral parts. Description
of leaf morphology follows Hickey (1984), trichome description follows Theobald
et al. (1984) and basic cell and tissue terminology follow Metcalfe (1984).

Habit and Architecture

The majority of Cybzanthus species in Ecuador and Peru are large shrubs
or small trees to 18 meters tall. Four species are small shrubs or subshrubs
(Cybianthus croatit, C. humilis, C. lineatus, C. nanayensis) usually under one
meter tall. With the exception of two species in subgenus Grammadenia, all
species are terrestrial. Cybianthus marginatus is an obligate epiphyte, while
C. magnus is a facultative epiphyte, frequently found on large tree trunks,
but also known from large rock outcrops.

Most members of Cybianthus exhibit Rauh’s Model of architecture (Hallé
et al. 1978), characterized by a polyaxial, monopodial, rhythmically grow-
ing, readily distinguishable trunk, that develops tiers of branches morpho-
genetically identical to itself. All branches are orthotropic and monopo-
dial, with lateral Keceaae inflorescences that do not affect shoot development.
However, two subgenera (Comomyrsine and Triadophora) exhibit Corner’s Model
(Hallé et al. 1978), characterized by a single aerial meristem that produces
a monoaxial (unbranched) axis on which inflorescences are lateral (axillary).
The resulting monocaulous tree is pleionanthic (not hapaxanthic, or mono-
carpic) and growth is indeterminate. The trunk may grow rhythmically or
continuously. Sporadic occurences of this architectural model occur in members
of subgenus Wezgeltia, from the Guianas and eastern Amazonia, however,
none of those species are known from Ecuador and Peru.

ile conducting fieldwork in Peru, a collection of Cybianthus kayapit
(Lundell) Pipoly was observed reiterating. In the population observed at
Explorama Lodge near Yanamono, (Loreto, Peru) one individual (Pipoly et
al, 12383), exhibited bayonet reiteration, caused by destruction of the api-
cal meristem of the trunk. The individual bore a reduced staminate inflorescence
with flowers slightly larger and more maroon in color than normal, and smaller
leaves with shorter petioles. As I have noted elsewhere (Pipoly 1987, 1992a),
leaves on the reiterative shoots resemble those of saplings. Variation in
inflorescence and floral morphology seen in this Peruvian population is consistent
with variation reported in Pipoly (1992a). Therefore, it appears that changes
in sex expression are consequent to a reiteration phenomenon, rather than
circumstantially associated with it. I postulate that reiterative branches may
be hormonally juvenile and as such, ically incapable of producing
bisexual or pistillate flowers, and produce only sapling leaves until flowering
occurs. This process may account for the great variation in leaf morphology
and apparent sexual lability often associated with many species of Cybzanthus.
Unfortunately, for individuals of normally dioecious species with monoaxial

4 Sipa 18(1)

models, it is not possible to determine if sex change has occurred because
no inflorescences are usually observed before the original apical meristem
is destroyed, and no inflorescences on other branches are available for com-
parison. However, precociously flowering individuals of Cybranthus incognitus
(P. Barbour 2405) support the hypothesis. Clearly, pruning experiments 77
situ will be necessary to test this hypothesis.

Branchlets

A branchlet is here defined as the distal 10 cm of any branch. Branchlets
may be flexuous, or straight; terete or angular; smooth, verruculose, or ver-
rucose-papillate (Fig. 1A, 1B), glabrous, glandular-papillate (Figs. 1D, 3F,
AF), ferrugineous stipitate-lepidote (Fig. 2A—F), ferrugineous or rufous dendroid-
and/or stellate-tomentose (Fig. 3A), rufous sessile furfuraceous lepidote, with
a tomentum of malpighiaceous trichomes (Fig. 3D), glandular-granulose,
with hydropotes (Fig. 3C), epunctate or rubiginous punctate-lineate, bear-
ing lenticels or not.

Leaves

Species of Cybranthus, like all Myrsinaceae, are exstipulate and have simple
leaves. The leaves are usually alternate, but may be subopposite or
pseudoverticillate, especially in subgenera Microconomorpha, Comomyrsine, Wergeltia
and Cybianthus. In the majority of species, the leaves are petiolate, but in
subgenus Grammadenia they are sessile. The ptyxis (the form in which one
single leaf is folded on itself in bud) is most often supervoluce (Cullen1978),
but it has not been adequately studied thus far for each subgenus. The leaf
vernation sensu Cullen (1978), is at least analagous with floral aestivation,
or the relationship of one leaf to another; in Cybianthus, it is in fact, un-
known. The leaf blade texture may be membranaceous, chartaceous or co-
riaceous. The shape is most often elliptic, but may be oblanceolate, lan-
ceolate, obovate, rarely ovate or oblong (Cybianthus marginatus). The apex
may be acute, acuminate, long-attenuate, rounded, obtuse, or emarginate,
without mucro or rarely (subgenera Grammadenia, Comomyrsine, Triadophora)
mucronate (Fig. 21). The base is rarely auriculate (subgenus Grammaenia)
or obtuse approaching auriculate (Cybianthus grandezti, C. kayapil, C.
anthuriophyllus) and is usually cuneate, long-attenuate, acute or obtuse, decurrent
on the petiole or not. The adaxial surface may be nitid, pallid, or sordid,
rarely scrobiculate , pusticulate or pustulate when mature. The blade mar-
gin may be regular or irregular, flat inrolled or revolute, entire, or rarely
undulate, lobate, crenate or dentate (Cybianthus pastensis), densely and mi-
nutely serrulate (C. anthuriophyllus), or coarsely dentate (C. schlimii, some
populations of C. pastensis, C. poeppigii, ), or with a very subtle but vascular-
ized vein ending (C. verticilloides) opaque, or rarely scarious (all species in

Pipoty, Cybianthus in Ecuador and Peru 5

Fic. 1. SEM photos of morphological features in sub G tenia. A. Cybianthus marginatus
Sees 6), showing verrucose papillae. B. Ge close-up. C, D.C. lineatus (Pipoly et al.

showing stem surface. D. Glandular papillae. E. C. costaricanus subsp. morii a
Haas pollen, equatorial and polar views. (Bars in figs. equal: A. 50 pm, B. 22.
48 yim, E. 12.3 pm) Figure reproduced from Pipoly, 1987.

)

6 Sipa 18(1)

Fic. 2. SEM ie one i ipi ( | Conomorpha

A. Cybianthus iolotype). B.C. eee aan C.C, irs 10803).
DG: eee ene 2821). E. C. Wien ianus (Killip &. Smith 26073). F.C. guyanensis
subsp. psewdoicacoreus (Simpson & Schunke 784). (Bars in Figs. A-F equal: 50 pm).

Cybianthus subg Grammadenia, Cybianthus himilis, and C. croatii of Cybianthus
subgenus Comomyrsine).

Leaf punctation and punctate-lineation may be pellucid (translucent), black,
orange red, or rubiginous (light red) in color. Punctations as defined by Pipoly
(1987) are definded as rounded, lyso-schizogenously formed cavities and
are distinguished from punctate-lineations, which are homologous, linear
cavities in the leaves. Among taxa bearing non-pellucid punctations or punctate-
lineations, all young, undexpanded leaves have orange glands, even if ma-
ture leaves have red or black glands. Experiments to determine integrity of
punctation color have revealed that it is not a reliable taxonomic character

PipoLy, Cybianchus in Ecuador and Peru Fi

NA; ph

Triado sis a

Fic. 3. SEM photos of represen I Laxiflorus
and Comomyrsine. A. Cy fe piule opulver, wena subsp. maonilepebis (Cowan & Soders

2146), branchlet, nee stellate trichomes. B.C, spicatis (Maguire & Politi 28004), ee
hydropote in abaxial lea _C.C. pastensis (Killip & Garcta 33886), branchlet, showing
developing nee ss c schlimti (Fosberg 20148), branchlet, showing malpighiaceous
trichome. E. C. kayapii (holotype) Staminate flower, showing vestigial pistillode, structure
a sai sera EC. humilis ae Jj.v.), showing g eae papillae of abaxial leaf sur-

ce. (Bars in Figs. equal: A-C. 50 pm, D & EF. 25 pm,

(Pipoly 1987). The other punctation character states include “conspicuous”
versus “inconspicuous,” reflecting the ease of viewing the punctation (a reflection
of its relative diameter in transverse section), and “prominent” versus “not
prominent,” which refer to whether the punctation is blister-like or planar,
respectively.

The abaxial and/or adaxial surface of the blade may be glabrous, or with

8 Sipa 18(1)

a vestiture composed of stipitate ferrugineous lepidote scales (subgenus
Conomorpha, Fig. 2A-2F), sessile furfuraceous lepidote scales (subgenus We7ge/tia),
dendroid and/or stellate trichomes, translucent glandular-lepidote scales,
or with scattered glandular papillae (subgenus Comomyrsine, Figs. 3F, 4F ),
malpighiaceus trichomes (subgenus Triadophora, Fig. 3D), or hydropotes
(subgenera Laxiflorus, Microconomorpha, Grammadenia, Comomyrsine, Figs. 3A,
3C, 4A-E ). The adaxial surface is usually glabrescent.

Hydropotes (“water drinkers”) were previously thought to be unique to
subgenus Grammdenia (Pipoly 1987), but have now been found in al
cies of subgenus Comomyrsine. Described by Mayr (1915) and Griiss (1927a,
1927b), hydropotes have been reported for numerous submerged aquatic
angiosperms (Fahn 1979; Gessner & Volz 1951). Elegance ultrastructural
and autoradiographic studies undertaken by Liittge (1964) and Liittge and
Krapf (1972) demonstrated the mineral absorptive function of hydropotes.
While both subgenera bear these structures, their morphology is strikingly
different. The hydropotes of subgenus Grammdenia (Fig. 5) consist of five
to seven subsidiary cells, a central foot cell, a basal cell, a stalk cell, and up

—_—

spe-

to 12 cells forming a scale-like cap, while those of subgenus Comomyrsine
consist of five to seven subsidiary cells, a central foot cell, a stalk cell, and
up to 12 cells forming a lenticular cap or globose body. In subgenus Comomyrsine,
the cap cell is formed before elongation of the stalk cell. In Cybanthus vertictlloides
(Fig. 4B), C. sprucei (Fig. 4D), and C. kayapii (Fig. 4E) the cap consists of a
spherical body of cells, while in C. croatii (Fig. 4A) and C. simplex (Fig. 4C),
the cap is lenticular. It is interesting to note that within subgenus Grammadenia,
the shape of the cap is the same among all species, while in Comomyrsine,
there are 4 types. The occurrence of hydropotes in subgenus Laxéflorus, in
pits over the abaxial leaf surface, and in subgenus Microconomorpha, ander
the papillate tomentum of the branchlets, was unknown before the present
study. Despite these differences, the structures are homologous, based on
position, function and ontogeny. In early ontogeny, a mucilaginous sub-
stance accumulates around the base cell (Fig. 7E). Later, the cap breaks off
(Fig. 7F), leaving a mucilaginous ring around the broken basal cell (Pipoly
1987, unpubl. data).

Subepidermal fibers, visible as numerous, thin, parallel lineate structures

on both surfaces of leaf blades in subgenus Triadophora are unique to it. They
have recently been shown to be homologous with those of C/avija in the
Theophrastaceae (Pipoly, unpubl.).

Petioles are present in most taxa, with the notable exception of subge-
nus Grammdenia, where the leaves are sessile. The petioles may be canaliculate,
marginate, or canaliculate with margins, trigonal, or rarely subterete, and
may be pulvinate (abruptly swollen basally) or gradually tapering basipetally.

Piroty, Cybianthus in Ecuador and Peru 2

Fic. 4. Representative SEM EM photos of morphological features in aan Comomyrsine. A—
E, ca ue of abaxial leaf surface. A. Cyhbianthus croatii paencie, showing lenticu

ipuBaG, gia (holotype), showing globose cap. C. C. simplex (Zak 1350), oon
ae cap. D vs aie Cees 15743), showing pigbese cap. E. C. kayapii (holo-
type), sh | I renee EC. kayapii (holotype), branchlet,

lowine g landulas iia (Bars in ies equal: A. 25 pm. B. 20 pm. C. 10 pm. D-F, 2
yim).

Cataphylls and Pseudocataphylls

Cataphylls (Fig. 9B) and pseudocataphylls are synapomorphic to subgenera
Comomyrsine and Triadophora, respectively. Earlier (Pipoly 1987), I had mis-
takenly interpreted them as inflorescence bracts. They may be alternate or
pseudoverticillate, alternating with pseudoverticels of leaves, or apparently
axillary to them, rigid, chartaceous, or membranaceous, linear-subulate to

10 SipA 18(1)

jm 4

Fic. 5. Ontogeny of hydropotes in subgenus Grammadenia. A, B, & D. Cybianthus costaricanus
subsp. costaricanus (Pipoly 7608), SEM photos. A. Mature sine oh cap with at least 8
cells, and with 5 subsidiary cells. B. Younger hydropote, cap with 4 cells, and with 6 sub-
sidiary cells. C. C. costaricanus eae mMOril 0g 7017), pee tion of young hydropote,
showing subsidiary cells (sc), f oe (fc), stalk cell (st), cap cell (ca), basal cell not dis-
cernible. D. Young hydropote, with 2-celled cap, showing aerate substance (ms).
E, F.C. costaricanus subsp. nee ao 7056), paradermal sections. E. Showing ma-
ture ai with 1 2-celled-cap. F. Showing hy ea after . has broken off. (Bars
in Figs. A-F equal: A & C. 28 pm, B. 20 pm, D.

ced from Pipoly, 1987.

m, E. 36 pm, F 53 pm. Figure

reproc

Pipoty, Cybianchus in Ecuador and Peru 11

acicular, rigid to membranous, keeled or flat, apicay crema. or not,
prominently punctate or punctate-lineate, glabrous, t, glandular-
papillate, bearing hydropotes, or orange lepidote scales, sessile or on a peti-
ole to 2 mm long. Pending further developmental studies, [am distinguishing
cataphylls from pseudocataphylls based on the absence of a petiole in the
former and its presence in the latter. Within Cybianthus, pseudocataphylls
are unique to subgenus Triadophora, while cataphylls are unique to subge-
nus Comomyrsine. Both pseudocataphylls and cataphylls may be distinguished
from an inflorescence bract by the fact that neither of these structures are
axillant to an inflorescence, neither are ephemeral, but both occur in pseudowhorls.

Inflorescence and Flowers

The inflorescence in Cybianthus is always lateral (axillary), and it may consist
of asimple raceme (erect or lax), a subpyramidal or pyramidal panicle (sometimes
thyrsoid), a spike, or rarely, a pletochasium. At times, species with essen-
tially racemose inflorescences may produce a panicle consisting of a pseudoverticel
of racemes ona reduced peduncle. In this treatment, an inflorescence is considered
spicate if the pedicels are obsolete or less than 0.3 mm long and subspicate
if the pedicels are from 0.4—0.8 mm long. In most species, the staminate
inflorescence tends to be longer, slightly more lax, and bears greater num-
bers of flowers. In those species bearing panicles, the staminate ones bear
secondary branches of the inflorescence that are longer, more floriferous, and
at times, more branched than the pistillate and bisexual panicles.

The inflorescence bracts are early caducous and are rarely seen in the field
or on specimens. The secondary branch bracts of panicles are also rarely collected,
with the exceptions of subgenera Comomyrsine and Triadophora. The floral
bracts may be membranaceous or chartaceous and are usually persistent in
staminate inflorescences, but are at times caducous in pistillate ones. The
floral bracts may be glabrous or bear a tomentum either adaxially, abaxially,
or on both surfaces. The floral bract apices are acute, attenuate, or acicular,
the margins entire or undulate.

The pedicels are cylindrical, clavate, or obconic, and at times accrescent
or crassate in fruit. In most species, the pedicels are subtended by an axillant
floral bract, but in Cybianthus kayapii, it is inserted at about the middle of
the pedicel.

Figure 6 illustrates the tremendous variation in floral morphology among
members of the genus, along with representative staminate and pistillate
flowers from Embelia, the paleotropical sister genus of Cybianthus. The flowers
may be functionally staminate, pistillate or bisexual. Consequently, the plants
are normally functionally dioecious, but may also be bisexual or polyga-
mous. Flowers are normally perfect, but in some species of subgenera Wezge/tia,
Comomyrsine and Cybianthus, the pistillode is often obsolete in the stami-

12 Sipa L8(1)

Fic. 6. Representative flowers of Cybianthus subgenera and sister genus, Embelia, open flowers
in anthesis above, calyx lobes below. A-B, Subgenus Microconomorpha, (Cybianthus pastensis),
note monomorphic flowers. A. Staminate flower (Ki//ip & Garcfa 33886). B. Pistillate flower

Prpoty, Cybianthus in Ecuador and Peru 13

nate flowers. The staminate flowers are usually larger than the pistillate ones.
The flowers are usually 4 or 5(—G)-merous, but they are 3-merous in subge-
nus Triadophora (Fig. 6K, 6L). The majority of species are homomerous, but
Cybianthus Rayapii has heteromerous flowers, (the calyx 4- and the corolla
5-merous), and in C. anthuriophyllus, the calyx 6- and the corolla 5-merous.
Either or both whorls of the perianth may be membranaceous, chartaceous,
coriaceous or carnose, epunctate, punctate or punctate-lineate. The punc-
tations may be inconspicuous or conspicuous, prominent (raised) or not,
pellucid, brown, orange or red. The calyx may be valvate or imbricate, and
may be cotyliform, cupuliform or urceolate. The corolla is valvate or im-
bricate, and may be campanulate, cupuliform, tubiform, infundibuliform,
salverform, rotate or subrotate. The stamens and staminodes are similar in
morphology, but the staminodes are smaller. The filaments of the stamens
and staminodes are partially united at the base to form a conspicuous or
inconspicuous, membranaceous, chartaceous or carnose tube, except in subgenus
Cybianthus, which has a terete or rarely, angulate staminal tube develop-
mentally fused with the corolla tube, the stamens thus appearing epipetalous.
In some species, the tube bears lobes (sterile projections of tissue) which
alternate with the apically free portion of the filaments. The apically free
portions of the filaments may be terete, flat, or absent. The anthers may be
dorsifixed, basifixed or versatile. Anthers may be lanceolate, ovate, cuadrate,
or deltate, apiculate, acute, truncate or emarginate, the apiculate ones may
have the apiculum erect, distally or proximally recurved, glabrous or glan-
dular-papillate. Antherodes of the pistillate flowers are similar to the an-
thers but reduced insize, and normally devoid of pollen, but occasionally,
they may produce copious amounts of abortive pollen (Pipoly 1983a). The
connective may be epunctate or prominently punctate, or rarely glandular-
papillate. The pollen is tricolporate and psilate (Fig. 1E). The pistillode
may be lageniform, conic or obturbinate. The pistil may be obnapiform,

de 3226). C—-D, Subgenus Stap eee stapfi2). C. Staminate flower (Fassett 25706).
istillate flower a sas 122 F, Subgenus Laxiflorus Sci Spicatus). E.
minate flower (Maguwire 35525). F. Cae flower (Prance et al. 3359). G-H, Subgenus
Conomorpha Cia. Se G. Staminate flower (Gareéa-Barri ga E Jaranill 19841).
H. Pistillate flower oe et al. 744). IJ, Subgenus Comomyrsine. 1.8 Cybianthus
sprucei (Cuatrecasas 16272). J. Pistillate flower etal kayapii (Acosta-Solis 5544), K-L,
as iene eee schlimii). K. Staminate flower (Haught 1502). L. Pistil-
late flower (Herrera 172 ie Eben Mies, 2 bianthus sp. nov. M. Staminate flower
(Matas & Weira 4456 )).N illate A awe 40590). O. Subgenus citleaes
ea flower, (Cybianthus sintenisii, P. Wilson S. a P. Subgenus Grammadenia, bis
wer, ee piresii, Maguire et al. 37052). Q-R, a Cybianthus, (Cybianthus
goyazensis). Q. Staminate flower Urwin & Soderstrom 5378). R. Pistillate flower (Herringer
45). $-T. ae sp. S. Staminate flower (Petefot 3887). T. Pistillaee flower (C. Wang 76408).
Drawing prepared by Peggy Duke.

4

14 Sipa 18(1)

conic, ellipsoid, or obturbinate, with a punctiform, or capitate stigma. The
stigma is large and capitate, with erose-fimbriate lobes and is caducous in
subgenus Comomyrsine. The ovary in species from Ecuador and Peru are ter-
ete. The placenta may be coryliform, cupuliform or globose, with 24 uniseriate
ovules naked or partially immersed. The fruit is a globose or depressed-globose
drupe and is one-seeded, with a thick or thin exocarp.

ANATOMY

This treatment is not intended to serve as a monograph of the genus, but
a few salient anatomical features may prove useful in identifying sterile material
or wood samples. Druses (Pipoly 1987-Fig. 8B—C) are ubiquitous in Myrsinaceae,
as are pericyclic fibers (Pipoly 1987, Fig. 8A). All Myrsinaceae have resin
ducts (canals) at least in the cortex, and in the field, copious amounts of
resin are visible in the canals of the pith and secondary phloem, in freshly
cut branchlets, in species of subgenera Grammadenia and Laxiflorus. Aeren-
chyma in the cortex of the primary stem is unique to subgenus Grammdenia
and is found in all species (Fig. 7F), except in Cybranthus lineatus (Fig. 7E).
The pith in primary stems of subgenus Wezgeltia is parenchymatous with
large, rounded starch grains, while that of Cybianthus magnus has angular
collenchyma (Pipoly 1987). It is interesting to note that the collencyma of
the outer cortex in Cybianthus magnus subsp. asymmetricus is tangential rather
than angular, while that of C. magnus subsp. magnus is angular (Fig. 8 D—
F). All species of Cybianthus have cortical vascular bundles, that may be
amphicribal (Fig. 7C) or hemiamphicribal (Fig. 7D), with (Pig. 7C) or without
(Fig. 7D) accompanying perivascular fibers. Wood of subgenus Comomyrsine
is notable for its thin-walled vessels (Fig. 9D), while the fiber-tracheids of
subgenus Grammadenia have walls so thick as to significantly occlude the
lumina (Pipoly 1987- Fig. 7C). Also, starch is present in the phloem fibers
of subgenus Comomyrsine (Fig. 9E, 9F) but not in Grammdenia. In leaf anatomy,
ic is notable that subgenus Grammadenia has functionally acrodromous ve-
nation. This is due to its unique leaf-node continuum, and consequent pri-
mary vascular system (Pipoly 1987-Figs. 7-9; 11-12) where the cathodic
and anodic leaf traces are autonomous from each other and from the rel-
evant midrib trace, a system thus far unique among angiosperms. Leaves of
subgenus Triadophora may be easily recognized by their subepidermal fibers,
while those of Cybianthus lineatus are unique for their bifacial palisade layer
(Pipoly 1987- Fig. 14b).

ECOLOGY

In Ecuador and Peru, species of Cybianthus occur in wet or moist, tall
terra firme forest on laterite, limestone or white sand, seasonally inundated
igapo or varzea, premontane humid, wet or pluvial forest on laterite or sandstone,

Pipoty, Cybianthus in Ecuador and Peru 15

a ‘
an ON ewegene
7

Fic. 7. Primary stem histological features in smpeencs lenta. A-B. Cybianthus ptariensi
(Pipoly et al. 7133), periderm formation | cork devel vee Ger Gree; mearginatns
(Pipoly 6546), amphicribal corical bundle Giecont ee iscular fibers. D. C. eile (Pipoly

et al. 7133), h emiamphicribal bundle with perivascular fibers (pf). E. C. dimeatus (Pipoly
7229), section showing parenchymatous pith, inner and outer cortex. F.C. Te

6546), section showing eee inner cortex; ae enchymatous pith and outer cor-
tex. (Bars in Figs. A—F equal to: A. 30 pm, B. 120 pm, C. 48 pm, D. 60 pm, E. 465 pm,
F. 120 pm. ae ee ce Bisoly. 1987.

cloud forest, elfin forest, montane or subpdramo thickets, or sandstone scrub
at high elevation.

The terra firme and premontane forest life zones are the richest in endemics,
with six and five species, respectively. Terra firme is here divided into two

16 Sipa 18(1)

Fic. 8. Sees stem histological features of subgenus Grammadenia. A.C. ns
morii (Pipoly 7017), showing pericyclic fibers (arrow). B. C. castaricanus subsp. costaricanus
(Pipoly 7068), showing druses in pith. C. C. costaricanus subsp. panamensis (Pipoly 7056),
showing druses in pith, using a light. D, E. C. magnus subsp.magnus (Pipoly 6453),
showing angular pith collenchyma. E. C. magnus subsp. asymmetricus (Luteyn, Pipoly et al.
10415), ee transitional srangent collenchyma. (Bars in Figs. ALE F equal to: A

195 pm, B. 45 pm, C. 48 pm, D. 230 pm, E. 74 pm, F. 60 pm.) Figure reproduced from
Pipoly, 1987.

edaphic categories, lateritic and white sand (often referred to as “varillal”
by Peruvians) and is defined as lowland tall moist or wet forest which is not
inundated. The canopy normally reaches 35 meters, with very few emer-

ent individuals. In the lateritic terra firme, Cybianthus kayapit, C. schlimii,
C. pseudolongifolius, C. vasquezii,C. cenepensis, C. grandezit, C. fuscus, C. jensonit,

Pipo.y, Cybianthus in Ecuador and Peru

tL}
,
O10,
@ @, y
eANS

Fic. 9. Morphological and anatomical
Pipoly et al, 12490). A. Habit, showing bayonet reiteration.
bipinnate panicle and leaf tapering to obtuse base. C. Stem apex, showing cata
Transverse wood section, showing thin-walled vessels. E. Tangential wood section, show-
ing rays and phloem fibers-tracheids. F. Phloem fiber-tracheids, showing starch grains. (Bars
in Figs. A-F, A. 12.6 cm, B. 2cm, C. 1 cm, D. 400 pm, E. 100 pm,

B. Habit, showing pyramidal,
hylls. D

18 Sipa 18(1)

and C. venezuelanus are found. Cybianthus kayapit, C. schlimii, and C. grandezii
are found on steep banks of small creekbeds, C. cenepensis and C. venexuelanus
are ridgetop species, and C. cenepensis, C. jensonii, C. vasquezii and C. psendolongifolins
grow along terraces above larger rivers. Vasquez (1997) has described “Irapayal,”

“Yarinal” and “Supay Chacras” associations within the lateritic serra firme, but
with so few comparative, quantitative forest inventories completed for both
countries, Iam not able to precisely list the Cybianthus species known from
each. The ferra firme forests on white sand (varillal) are shorter in stature than
those on lateritic soils, and generally support lower numbers of lianas. Amon g
the varillal forest types described for Peru, Vasquez (1997) lists “varillal seco,’

“varillal hamedo” and “chamizal” or “ojo de varillal” associations, only some
of which have been noted on label data. However, Cybianthus peruvianus, C.
spichiger1, and C. gigantophyllus occur on rolling hills in the varillal forest
type. Among those species which occur on steep hillsides near light gaps,
Cybianthus gigantopyllus is most notable, as it occurs in the ecotonal area
between forest gaps and mature forest. Cybianthus nanayensis, a subshrub, i
frequently found in gaps left by large treefalls in overmature forests, ca
along trailsides, where it occurs in the rather dense herbaceous stratum. Cybianthus
restnosus, another inhabitant of the terra firme forest on white sand, occurs on
terraces above black water rivers in the forest, while C. nestorii is found in
the more open shrubby “varillal seco” transitional area near the riverbank.
Cybranthus fulvopulverulentus sabsp. magnoliifolius typically occurs in campina,
or Campinarana formations i in Brazil, but in Peru it has been collected once
in the “varillal seco” an open shughie area on white sand several hundred
meters from a black water riverbank. Unfortunately, no literature directly
addressing this forest type is known for Ecuador.

The two forest types subject to inundation have been divided into varzea,
flooded by white water, and igapé, flooded by black or black and white water.
The other significant Je eae between these two forest type is that in varzea,

the forest is indundated for a much shorter time than that of the igap6. To
date, I know of no documentation for occurrences of forests inundated by
clear water in Ecuador or Peru, as they are found elsewhere in Amazonia
(Brazil, Colombia, Venezuela), but are best developed in Colombia. Three
species are known from igap6, with no endemics. Cybianthus giyanensis subsp.
pseudorcacoreus is found in the igap6 at its limit with terra firme, while C.
penduliflorus is found well within the igap6 and is frequently found in standing
water. Cybianthus spicatus is found both in igap6é and varzea, and exhibits
both staminate and pistillate ecotypes, with some exhibiting apparent ran-
dom variation. It is a broadly ranging polymorphic ochlospecies (sensu Pipoly
1983a) with great morphological variation. The known Peruvian popula-
tions, from Hudnuco and San Martin, are identical to those found in cen-
tral Guyana (Pipoly 198 3a). The other varzea species, Cybianthus cyclopetalus,

Pipoty, Cybianthus in Ecuador and Peru ee

is known only from Madre de Dios in Peru, and from the Jurua area in Brazil.
It grows in the margin of varzea near its junction with the serra firme forest.
Forest associations within the vazea have been described by Vasquez (1997),
but I have been unable to match the corresponding complement of Cybianthus
species, owing to inadequate label data. The three forest associations present
in varzea for Peru include, “barrillal,” “restinga” and “bajial.” Clearly, more
fieldwork is needed to discern foristic differences among these association
types.

Premontane forest habitats are found from scarcely above 200 m to nearly
1,000 m. The forest is lower in stature than the serra firme, have a greater
epiphyte load, and a larger number of lianas. Those on sandstone are dis-
tinguished here from those on other soils. On sandstone, three species are
found of which one, Cybianthus timanae, is endemic. The other species, Cybianthus
comperuvianus, a new taxon described herein, is known from these forests in
Peru, Bolivia and adjacent Brazil. The other premontane forest formation
on lateritic soils hosts 9 species, Cybsanthus minutiflorus, C. huampamiensis,
C. granulosus, C. poeppigii, C. schlimit, C. peruvianus, C. fosteri, and C. venezuelanus
and C. flavovirens. Among these, only Cybianthus venezuelanus, C. poeppigit,
C. schlimii and C. peruvianus are not endemic. A surprising new distribu-
tion record for Cybzanthus lepidotus, from Bagua Province, Imaza District,
of Amazonas Department, Peru, is recorded here. Cybianthus lepidotus was
once thought to be a Guayana Higland endemic, but was found in Bolivia
in the Maipiri region on the sandstone “laja” formations there (Pipoly 1992a).
This kind of disjunction, concomitant with those for species such as Cybianthus
spicatus, and Cybianthus lineatus (see below), support recent thematic map
data at NASA, showing that the Ecuadorean/Peruvian area north, slightly
east, and immediately south of the Cordillera del Condor, contains significant
sandstone formations that may constitute “tepui satellites” (sensu Maguire
1979). Clearly, much more exploration in southeastern Ecuador and north-
ern Peru, should be a high research priority.

There are six species of Cybianthus known from cloud forests. Cybianthus
pastensis and C. incognitus are found in areas of high shade and moisture within
these forests. Along the margins to the leeward side, in that portion of the
Chocé Floristic Province that extends into Ecuador, Cybianthus cuatrecasasi1
may be found, while on the eastern Andean slopes of Peru, C. /aetus grows
in exposed, wind-swept margins of the forest. The lack of records for Cybzanthus
/aetus in Ecuador is more likely a collection artifact than a reflection of its
rarity, given its occurrence in the Department of Boyaca, Colombia. It is
notable that Cybianthus patensis and C. incognitus, when growing at eleva-
tions below 1,000 m, are ridgetop species. Cybianthus magnus subsp. magnus
is an obligate epiphyte in closed cloud forests, growing in the forks of trees
or on deep detritis, and shielded from winds.

20 Sipa 18(1)

Elfin forests and montane thickets are transitional formations below the
jalca (a formation like a paramo but without species of the Asteraceae subtribe
Espletiniae). Elfin forests host considerable numbers of trees and usually
have trees up to 5 m tall. The montane thicket (subpéramo) grows in more
exposed areas at higher elevations and is dominated by low shrubs and small
trees to 2 m, with broad crowns. Cybianthus marginatus is found in both of
these habitats, but the leaves and stature of the plants are much smaller in
the thicket formation. Also, the verrucose papillae of the stem are much
more pronounced in the thicket habitats than in plants growing in the elfin
forest. Cybianthus magnus subsp. asymmetricus grows in open montane for-
ests or subparamo thickets, and elfin forests, where it is subject to exposure
to the high winds and rain. Its leaf variation is significant, but it is easily
recognized from subspecies magnus by the hydropotes of the adaxial leaf surface
and orange punctations of leaf, inflorescence and perianth parts and the unique
white, then lavender fruits.

Finally, sandstone scrub, called “pajonal,” is known thus far only from
Peru. It is the formation growing at the highest elevations where Myrsinaceae
occur, mostly well over 3,000 m. In these habitats, there are few, small shrubs
which rarely exceed 1.5 m tall. Cybtanthus lineatus is found in this habitat,
the first locality for this species outside the contiguous Guayana Highland.
No similar habitat has been described in Ecuador, but it may be present in
the Cordillera del Céndor.

TAXONOMIC CONCEPTS, NOTES ON KEYS AND SPECIMEN CITATIONS

My species concept follows that of Wiley (1978, 1981), who defined a spe-
cies as follows: “An evolutionary species is a single lineage of ancestor-descen-
dant populations which maintains its identity from other such lineages and
which has its own evolutionary tendencies and historical fate.” My subspecies
concept (Pipoly 1987), defines a subspecies as follows: “groups of populations
within a single lineage of ancestor-descendant populations that show varia-
tion by unique combinations of plesiomorphies, or homoplasic apomorphies,

correlated with biogeography and/or ecology. This rank is primarily used to
convey information regarding variation in the life histories of these populations
and character state differences hypothesized to be the result of this variation.
The subspecific rank in no way attempts to predict speciation events.”
The keys are artificial and designed to expedite identification of herbarium
specimens. An attempt has been made to emphasize vegetative characters to
increase the keys’ usefulness with sterile material. The numbers appearing be-
fore the taxa refer to their respective position in the key; any correlations with
phylogenetic relationships are coincidental. Quantitative and qualitative
data presented in keys and descriptions for floral parts and bracts were taken
from organs rehydrated from herbarium specimens by boiling in water.

Pipoty, Cybianthus in Ecuador and Peru 2

Measurements from these range from 10% to 15% greater than those mea-
surements taken directly from dried material. Data regarding stem diam-
eters, inflorescence rachises, pedicels, leaf, and fruit shape were taken from
dried herbarium specimens. Extra-Ecuadorean and -Peruvian specimens are
cited for all new species and for recent collections of other species used to
significantly amplify previously published morphological descriptions (Pipoly
1981, 1983a, 1983b, 1987, 1988, 1991, 1992a, 1993, 1994, 1995, 1996).

A description of the genus Cybianthus and a key to its subgenera in Jcua-
dor and Peru are provided below. This description, along with that of the
subgenera and species that follow include features found in each taxon as a
whole, including those populations and species occurring outside Ecuador
and Peru. Phylogenetic studies applicable to species in this treatment may
be found in Pipoly (1987) and in the forthcoming Flora Neotropica treatment.

TAXONOMIC TREATMENT
Cybianthus Mart., Nov. Gen. Sp. Pl. 3:87. 1831. nom. et typus cons.,G. Agostini,
Acta Biol. Venez. 10:141. 1980.; Pipoly, Mem. New York Bot. Gard. 43:46. 1987;
Pipoly, Ann. Missouri Bot. Gard. 79:913. 1992. Type species: C. penduliflorus Martius.
Terrestrial or epiphytic, monoaxial or polyaxial, dioecious, monoecious
or polygamous shrubs or trees to 15 m tall. Roots positively geotropic or dia-
geotropic. Branchlets glabrous, glandular-granulose, dendroid- and stellate-
tomentose, furfuraceous- or ferrugineous-stipitate-lepidote. Leaves sessile or
petiolate, alternate, subopposite, or pseudoverticillate, the venation
camptodromous or rarely acrodromous; petioles obsolete or when present,
canaliculate or marginate, tapering gradually to the base, or abruptly swol-
len toward the base, here termed “pulvinate.” Imflorescence: staminate, pistil-
late, bisexual or polygamous, lateral (axillary), a simple raceme, panicle of
racemose or spicate (rarely corymbose) branches, a pleiochasium, or an 1n-
determinate umbel appearing racemose. Flowers functionally unisexual or
bisexual, 3—G(—7)-merous; calyx cotyliform to cupuliform, the lobes im-
bricate, valvate or aberrantly contorted, basally connate 1/5—2/3 their length,
abaxially glabrous, glandular-granulose, ferrugineous stipitate-lepidote, or
translucent-lepidote, adaxially glabrous, epunctate or prominently orange,
red or black punctate, the lobes entire to erose-fimbriate, glabrous or glan-
dular-ciliate; corolla rotate, subrotate, cupuliform or campanulate, rarely
infundibuliform or salverform, the lobes imbricate or valvate, basally con-
nate 1/5—3/4 their length, abaxially glabrous, glandular-granulose, or fer-
rugineous stipitate-lepidote, adaxially glandular-granulose at least at the
junction of the tube and lobe, the margin entire to erose-denticulate, gla-
brous, glandular-granulose or rufous glandular-papillate; st d staminodes
adnate to corolla tube at least 2/3 their (enone the filaments variously con-
nate to form a tube, the staminal tube adnate to the corolla tube or at times

22 Sipa 18(1)

developmentally fused with it (thus the stamens appearing epipetalous),
bearing fleshy lobes alternate with the apically free portions of the filaments
or not, the anthers erect or distally curved, ovate, widely ovate, or triangu-
lar, basifixed or dorsifixed, apically acute, rounded, truncate, emarginate or
minutely apiculate, the apiculum erect, proximally or distally curved, ba-
sally truncate, cordate, or rarely hastate, deshiscent by pay pores, confluent
apical pores (birimose), or by wide or narrow longitudinal slits; pollen tricolporate,
psilate; staminodes morophologically similar to the stamens but greatly reduced
in size, the antherodes at times producing abortive pollen; pistl obnapiform,
ellipsoid, umbonate or obturbinate, the ovary sparsely to densely translu-
cent glandular-lepidote, the style glabrous, the stigma capitate, capitate-
lobate, or punctiform, persistent or early caducous, the placenta free-cen-
tral, carnose, umbonate or globose, the ovules campylotropous (1—)2—5(—7),
uni- or biseriate; pistillode conic, lageniform, obturbinate or irregularly shaped
vestigial pistillode, the pistillode hollow or bearing a sterile placenta, rarely
absent. Fruit drupaceous, 1(—2)-seeded, the endosperm translucent, non-
starchy, the embryo small, linear, flexuous, erect or curved, longitudinal or
transverse, the cotyledons not well-developed.

Distribution.—One hundred sixty-seven species; Nicaragua, Costa Rica
south through Panama to the Andes southward to Bolivia, from Colombia
eastward across Venezuela and Brazil and the Guianas, then southeastward
to the Atlantic coastal forests of SE Brazil.

Ecology.—Members of Cybianthus are principally riparian, occurring only
in primary forests or rarely in somewhat disturbed ones, and thus, may serve
as indicators of environmental quality. Throughout the range of the genus,
its members are known from wet tepuf savannas, moist scrub, cloud and
elfin forests (including “ceja de selva”), subparamo thickets, montane, premontane,

pluvial, wet and moist forests, paramo, jalca, igap6, varzea, varillal, campinas,
campo rupestre, restinga, cerrado, and caatinga vegetation types. In Ecua-
dor and Peru, the majority of the species occur in lowland and premontane
forests at the junction of Hylaea and the eastern slopes of the Andean Cor-
dillera (see ECOLOGY section).

Cybianthus is most closely related to the paleotropical lianous genus Embelia
(Pipoly 1987), and cladistically defined by the unique glandular-granules at
the corolla lobe and tube junction. For practical purposes of identification, the
combination of lateral racemes or spikes, or racemose or spicate panicles, and
filaments which are shorter than the corolla, connate at least 1/4 their length,
and adnate to the corolla at least 1/3 its length, allows for easy recognition.

KEY TO SUBGENERA OF CYBIANTHUS IN ECUADOR AND PERU

1. Corolla cotyliform, cupuliform, campanulate, or rarely salverform; anthers
longer than wide, distally recurved, apically acute or minutely apiculate.

Pipoty, Cybianthus in Ecuador and Peru 29

2. Branchlets sous or glandular-granulose; anthers dehiscent i narro

longitudinal s I. Mi Pee aor
2. Branchlets ae tomentose or stipitate-lepidote; anthers dehis-

cent by wide longitudinal s

. Branchlets and calyx ne, adda stipitate-lepidote; abaxial ps

eae glabrous near margin I. Conomorpha
. Branchlets ferrugineous tomentose, calyx glandular-granulose or ae
on rarely ferrugineous tomentose; abaxial corolla surface eee
Ill

lar-granulose near margin . Laxiflorus
1. Corolla rotate to subrotate; ice wider than long, erect, apically rounded

to truncateor emargin
4. aioe spay re basally; anthers dorsifixed, longitudinally
dehisc
De Bones monoaxial; anthers erect, not versatile
6. Stem glandular papillate, at times with yop, ee Sie ylls
apically and at times, alternating with the leaves; lea ithout
subepidermal fibers; flowers 4- or 5 -merous; coal ieee epunctate
or inconspicuously pellucid or orange punctate, glandular-granulose
within; stigma large, capitate-lobate, early caducous, the lc
gins fimbriate V. Comomyrsine
6. Stem with malpighiaceous hairs, without cataphylls, but at times
with aborted leaves (pseudocataphylls); leaves with numerous paallel
ace fibers (most easily seen adaxially); flowers 3-merous;

Ila lobes prominently black punctate, maculate, glabrous within
except at lobe and tube junction, stigma small, capitate-lobate, per-

sistent, the lobe margins entire V. eee
5. Plants polyaxial; anthers versatile I. Weigeltia
4, Petioles obsolete or not abruptly swollen basally; anthers basifixed aie
dehiscent
7. Leaves sessile, apically mucronate, basally auriculate, the margins

scarious; staminal tube merely adnate to corolla tube........... VII. Grammadenia
7. Leaves ae apically acute, acuminate or caudate, basally acute,

attenuate or cuneate, the > staminal tube development
ally fused to corolla tube, he stamens thus appearing epipetalou
III. Cybianthus

I. Cybianthus subgenus Microconomorpha (Mez) G. Agostini, Acta Biol.
50. 1980; Pipoly, Wrightia 7:235. 1983. Conomorpha A. DC. subgenus
Microconomorpha Mez in Engl., Pflanzenr. [V. 236(Heft 9):251. 1902. Microconomorpha
ee ae Wrightia 5:349. i ie Type Species: Conomorpha verticillata Zahlbr.,
. Naturhist. Hofmus. 7:3. 1892, non Mez (1902). = Pi cee ane

es os Agostini (LECTOTYPE: Abosdal Acta Biol. Venez. 10:150. 1980)

Cybianthus Mart. subgenus Iteoides G. vee ae ek Acta Biol. Venez. 10:148.
he Type Species: Badula iteordes Benth., Pl. Har 17. 1896. Conomorpha iteoides

+

(Benth.) Mez in Engl., Pflanzenr. IV. onc kh 9): ae 1902. Type Species: Cybianthus
ee (Beat ) G. Agostini, Acta Biol. Venez. 10:149. 1980.
Terrestrial monoecious, dioecous or polygamous shrubs or small trees. Bark
smooth to slightly fissured, light brown, thin. Root positively geotropic.
Trunk distinguishable, leptocaulous, the growth dynamics following Rauh’s

24 Stipa 18(1)

Architectural Model (Hallé et al. 1978). Branchlets thin, terete, densely fer-
rugineous glandular-granulose, the granules often stipitate. Cataphylls and
pseudocataphylls absent. Leaves pseudoverticillate; blades petiolate, often with
translucent glandular lepidote scales. Lnflorescence a simple raceme or bipin-
nate panicle, staminate, piscillate, or polygamous, the peduncle 1-4 cm long,
densely glandular-granulose; inflorescence and floral bracts, perianth and
pistil bearing prominently raised red or black punctations; inflorescence bracts
large, often foliaceous and persistent; floral bracts linear-lanceolate, the margins
glandular-ciliate, caducous, the pedicels erect, accrescent in fruit. Stami-
nate, pistillate and bisexual flowers monomorphic (similar in shape), the staminate
the largest, the pistillate the smallest in size, white to yellowish-green, (4—)
5(—G6)-merous; calyx cotyliform, the lobes valvate, the margins densely glandular-
ciliate; corolla cotyliform to campanulate, the lobes imbricate, glabrous without
except glandular-granulose near the margin, glandular-granulose over the

entire surface within; stamens and staminodes with a conspicuous staminal
tube, the apically free portions one to three times longer than the anthers,
the anthers elongate-triangular to ovate, prominently curved distally, apically
obtuse to apiculate, basally cordate to hastate, dorsifixed 1/3 to more than
1/2 length from base, dehiscent by narrow longitudinal slits, the staminodes
producing abortive pollen grains; pistil in pistillate and bisexual flowers
obturbinate, the ovary densely translucent-lepidote, the style thick, trun-
cate, the style punctiform, the placenta umbonate, bearing 3—4 uniseriate
ovules immersed in placental tissue, but exposed apically by placental pores;
pistillode similar to piscil but reduced in size, hollow or bearing 2 abortive
ovules. Fruit drupaceous, |-seeded, the exocarp thin, prominently black punctate.
Subgenus Microconomorpha contains 5 species, of which one, Cybianthus
pastensis (Mez) G. Agostini, is known from Ecuador and Peru. J earlier indi-
cated that there was no evidence to support subgenus I[feosdes as a separate
entity (Pipoly 1987), so it is treated here in synonymy under subgenus
Microconomorpha.
‘ ae pastensis (Mez) G. Agostini es 3c), Acta Biol. Venez.
10:1 1980. Compe ae oe in Engl., Pllanzenr. [V. 236(Heft 9):252.
BIA.

1902. Ma ell | Wrightia 5: 349. 1977. Type: COLOM
Narino: “Pdramo sale Purugai, Prov. Fe Pasto,” 2,500 m, 1866 (stam. fl), J. Triana
2585 (LECTOTYPE (Pipoly 1983a): W; IsoLEcrorypEs: C, COL, G, P)

Myrsine verticillata C. Presl, Relig. Haenk. 2 oe 1835. Conomorpha verticillata (C. Presl)
z in Engl., Pflanzenr. IV. 236(Heft 9):2 1902, non Zahlbr. (1 oe Conomorpha
preslit JE. eae. Candollea 5:398. 1934. neeenaan verticillata (C. Presl) Lundell,
Wrightia 5:349. 1977. Type: PERU. HuAnuco: without further poe Seat: date,

, 1. Haenke 98 (LEC TOTYPE (Pipoly 1983a): PR; ISOLECTOTYPES , W). Non

~~

(stam. fl.
Cybia sis verticillatus (Vell.) G. Agostini, Acta Biol. Venez. 10: 168. 1980.
pas verticillata Zahlbr., Ann. K. K. aghee Hofmus. 7:3. 1892, non C. Presl

PipoLy, Cybianthus in Ecuador and Peru 25

(1835) Conomorpha jelskit Mez in ae Pflanzenr. IV. 236(Heft 9):251. 1902, nom.

superfl. Conomorpha yee J.E. Macbr. jelskii (Mez) J.F. Macbr., Field Mus. Nat.

Mise. -Botsereka 201s O00. Me icieenorphe ee (Mez) Lundell, Wrightia 5:349.

1977, Type: PERU. cee A: Cutervo, Apr. 1879 (stam. fl.), C. von Jelski 11 (HO-
E: W; photo and fragment, F, F Neg. 31980).

Ge dentata Mez in Engl., PAlanzenr. IV. 236(Heft 9):252. 1902. Microconomorpha
dentata (Mez) Lundell, Wrightia 5:349. 1977. Type: ECUADOR. PicHINcna: In cor-
dillera from Quito to Tungurahua, 2,000—3,000 m, 1857-9 (stam. & bisex. fl.), R.
ee 5175 (LECTOTYPE a 1983a): K; isotecrotyPes: BM, BP, C, CGE, GH, GOET,
LD, LE, F Neg. 2

G ee Sa Mer in Engl., Pflanzenr. IV. 236(Heft 9):253. 1902. ne ead ip
ee oa Lundell, Wrightia 5:349. 1977. Type: PERU: without locality, wit

date (sta HJ. ae von s.n. (LECTOTYPE (Pipoly 1983a): G
Conomorpha panam indell, Wrightia 5:290. 1976. Micraconomorpha ee ee
Lundell Wrgha5 eS 1977. Cybianthus morii G. Agostini, Acta Biol. Vene 154.
1980. PANAMA. Cuirigut: Cerro Pando, on continental divide a oe

Costa a Kordee a. 16 km W of Hato del Volcan, 2,000—2,482 m, 20 Jul 1975
(stam. fl.), S. Mor: G A. Bolten 7292 (HOLOTYPE: LL-TEX; Isorype: MO).
Shrub or tree to 6 m tall. Branchlets and inflorescence densely ferrugineous
glandular-papillose, the branchlets angulate to prominently ridged, (1 .5—)
iam. Leaves pseudoverticillate; blades membranaceous to coria-
ceous, narrowly oblanceolate to oblong or obovate, (3.0—)6.5—14.5(—21.0)
cm long, (1.2—)2—4.5(-6.8) cm wide, apically attenutate, acute or acumi-
nate, basally cuneate, not decurrent on the petiole, prominently punctate
and minutely ferrugineous stipitate-papillose above and below, the midrib
impressed above, raised below, the secondary veins 7—15 pairs, prominently
raised below, the margin undulate, lobate, crenate or dentate, rarely subentire;
petioles marginate, (0.2—)0.5—2(—2.7) cm long, densely glandular-papillose.
Staminate, pistillate or polygamous inflorescence. monomorphic, erect or lax, a
simple raceme, 1.8—5.5 cm long, the rachis thin to thick, densely glandu-
lar-papillose; inflorescence bracts chartaceous, obovate to elliptic, (4.3—)6—
11 mm long, 3—7 mm wide, apically acute to acuminate, basally cuneate,
densely and prominently red punctate; floral bracts chartaceous, linear-lan-
ceolate, (0.8—)1.4—2.2(—7) mm long, 0.6—-0.8 mm wide, apically attenuate,
caducous; pedicels cylindrical, (1.3—)2—7(—7.5) mm long, densely glandu-
lar-papillose. F/owers (4—)S-merous, white to yellowish-green; calyx chartaceous,
shallowly cotyliform, (0.6—)0.8—1.1(-1.5) mm long, unequally divided, the
tube 0.2—0.5 mm long, the lobes suborbicular to very widely ovate, (0.4—)
0.6—1 mm long and wide, rounded to acute apically, glabrous, densely and
very prominently orange or black punctate, the margin subentire to erose-
dentate, densely glandular-ciliate; corolla chartaceous, cotyliform, 2—2.6(—
3.6) mm long, the tube 0.2—0.3 mm long, the lobes ovate to narrowly ovate,
1.7—2.3(—2.8) mm long, 0.8—1.3(-1.5) mm wide, highly reflexed at anthesis,
apically rounded to obtuse, prominently orange or black punctate; stamens

26 Sipa 18(1)

and staminodes 1—1.6(—2.5) mm long, the staminal and staminodial tube
0.7—-1.8 mm long, the apically free portions of the filaments 0.3-0.7 mm
long, the anthers elongate-triangular, 0.6—-1.2 mm long, apically obtuse,
basally hastate, the connective red punctate ventrally and dorsally, dorsifixed
ca. 1/3 to slightly less than 1/2 from base; pistil and pistillode 1.2—1.8 mm
long, the ovary (0.6-)0.8—1 mm long, 1—-1.3 mm diam., densely translu-
cent glandular-lepidote, the style thick, 0.5—0.8 mm long, the stigma punctiform,
the pistillode hollow or containing one abortive ovule. Fruit globose, green,
then red, then black at maturity, 3-4 mm diam. when dried exocarp thin,
prominently pellucid punctate.

Distribution.—Costa Rica to Colombia, southward to Peru, from 1,500—
3,200 m elevation.

Ecology and conservation status. —C ybianthus pastensis is known from premontane
and montane pluvial and cloud forests, and at elfin forest margins. Popula-
tions in areas exposed to winds have more coriaceous leaves and shorter stature,
frequently as small as one meter in height. The wetter the habitat, the more
membranaceous the leaves become, and the longer the inflorescences. Fieldwork
in Colombia has shown that populations may contain six individuals per
hectare, and that the population rapidly dwindles in areas of disturbance.
Owing to population dynamics thus far observed, Cybianthus pastensis should
be considered threatened.

Etymology.—The specific epithet refers to the area from which the type speci-
men was collected, near the city of Pasto, Department of Narifio, Colombia.

Representative specimens examined. COLOMBIA. a ae Mpio. Urrao, Parque
Nacional Natural “Las Orquideas,” Vereda eae Permanent Premontane Rainforest In-
ventory Plot, right bank of Rio Calles, 06° 32' N, 76° 19' W, poe 00 m, 29 Nov

1993 (fr), J. Pipoly, A. Cogollo et al. 17322 (BRI I, COL, JAUM, MO), limits of Parque Las
Orquideas, left bank of Rio Calles, 1,450-1,500 m, 30 Nov 1993 (ster.), J. Pipoly, A. Cogollo
et al. 17376 (BRIT, JAUM, MO), near limit of Parque Las Orquideas, Alto de Palmitas,
ca. 1 km from INDERENA Cabafia Calles, 1,300—1,400 m, | Dec 1993 (ster.), J. Pipoly.
A. rae 17505, 17523 (BRIT, JAUM, MO), 2 Dec 1993 (fl bud), J. Pipoly, A. Cogallo
etal. 17534 (BRIT, COL, JAUM, MO) res Wt ce of Rio Calles, 1,350-1,450 m, 7 Dec

1993 (sia am. fl), J. Pipoly et al. 17881 (B L, JAUM, MO); Along trail to Finca La
Quince, above Urrao, 06° 30' N, 76° 10! 7 eras 2,800 m, 21 Nov 1988 (stam. fl), G.
oe et al, 13212 (BRIT, HUA, MO); Mpio. Frontino, Regién de Murrf, ca.

m Nutibara, 06° 40' N, 76° 20! W, 2,000 m, 9 Dec 1988 (pist. fl, fr), G. McPherson et al
oe (BRIT, HUA, MO). Norte de Santander: San Antonio, W of Cali, near summit of
ordil aie are Ca 1,9800- 2,350 m, 26 Feb—2 Mar 1939 (stam. fl), E. Killip GA. Garcia
33886 (A, 8S, US). ECUADOR. Azuay: Chiguinda, on E slopes of cordillera E of Sigsig,
03° 12'S . 3° 36) W. 1 ,600—1,800 m, ie (stam. fl), E Lehmann 5143 (K-2 sheets). Carchi:
Paramo de Achupallas, 01° 46'S, 78° 33' W, 2 fe 000 m, 1899 (stam. fl), FE Lehmann
6202 (K-2 sheets); From Prima Vera about 6 hrs. hike up Rio Gualchan Drainage to Nilo
Ortiz’ shelter, 00° 50! N, 77° 72' W, 1,930-2,200 m, 7-8 Jun 1993 (fl bud), /. eV
al. 55 (BRIT, MO, QCNE). Loja: Cerro Bangala, ca. 10 km E of Yangana, 2,500— :

18 Oct 1988 (pist. fl, fr), G. Harling 25313 (GB), (stam. fl), G. Harling 25334 een

2

ee,
x

Pipoty, Cybianthus in Ecuador and Peru ee

Loja, Carretera Loja~-Zamora, at high point, 03° 58'S, 79° 04' W, 2,400—2,600 m, 23 Dec
1991 (pist. fl), D. Rubio et al. 2252 (BRIT, MO, QCNE); Loja, 3,500 m, 1 Dec 1876 (stam.
fl), E. André 4551 (F, K, NY); Divide between Quebrada Jipir and E fork of Rio Zamora,
W slope of Cordillera de Zamora (El Céndor), 9 km E of Loja, 04° 00'S, 79° 06' W, 2,700
m, 19 Feb 1945 (stam. fl), ER. Fosberg G M. Giler 23119 (NY, US); Loma de Loro, 6 km §
of Saraguro, on Rd. to Loja, 3,200 m, 11 Feb 1985 (stam. fl), G. Harling & L. Andersson
21594 (AAU, S); Saraguro-Loja Rd., km 12.4, turnoff toward Fierro Urco, Km 3.
03° 42! 33"8, 79° 18'03" W, 3.120-3,391 )m, 7 Dec 1994 (pist. fl), P. Jorgensen et al. 1297
(BRIT, LOJA, QCA, QCNE); W slope of Nudo de Sabanilla, ca. 8 km above Yangana on
Rd. to Valladolid, 2,300—2,500 m, 2 Apr 1985 (bisex. fl, fr), G. Harling & L. Andersson
23540 (GB); Cerro Toledo, Rd. to La Torre, ca. 7 km SE of Yangana, 2,500 m, 7 Apr 1985
(stam. fl), G. Harling & L. Andersson 23842 (GB). Morona-Santiago: Between Campanas
and Arenillas, along Rio Tintas, 10 leagues SE of El Pan, 2,195 m, 13 Jul 1943 (stam. fl),
J. Steyermark 53642 (F, NY); Above Mirador, 2,375 m, 9 Sep 1943 (stam. fl), J. Sigermate
53897 (EF, NY). Napo: 10 km W of Cuyuja, along Quito-Lago Agrio Rd., 00° 25'S, 78
00! W, 2,700 m, 31 Apr 1983 (stam. fl), H. Balslev 4295 (AAU, QCA); Salcedo-Napo Rd.,
2,390—2,590 m, 7 Feb 1977 (fr), J. Brandbyge 42095 (AAU, QCA); E of Borja, Cerro Antisana,
28 Jul 1960 (stam. fl), P. Grubb et al. 1073 (OXF, NY); Sta. Barbara Scumbios, 00° 22'S,
77° 10! W, 2,700 m, 10-15 Feb 1959 (stam. fl), L. Holm- sea 6818 (AAU, QCA); 8-12
km ESE of Sta. Barbara, 00° 40' N, 77° 30! W, 2,780—2,880 m, 11 Jan 1985 (stam. fl), J.
Bi G E. Cotton 11011 (GB, QCA, NY, VEN); Paso de panei io Chalpi, at bridge
yn Papallacta-Baeza Rd.., 2,800 m, 6 May 1967 (stam. fl), B. Sparre (5046 (GB); Cantén
oe Sierra Azul (Agricola Industrial Rio Aragén), 00° 40! S, 77° 55' W, 2,300 m, 2
May 1992 (fr), A. Alvarez et al. 381 (BRIT, MO, QCNE), (fr), A. Alvarez et al. 412 (BRIT,
MO, QCNE), Campamento Estero Chico, 00° 41'S, 77° 56' W, 2,500 m, 18 Jun 1992
(fr), A. Alvarez et al. 490 (BRIT, MO, QCNE); Sierra Azul, Cordillera de Huacamayos, 00°
41'S, 77° 54! W, 2,500—2,700 m. 10 Feb 1994 (fr), A. Alvarez et al.1330 (BRIT, MO,
QCNE). Tungurahua: On Patate-Triunfo Rd., 01° 18'S, 78° 25' W, 2,950 m, 5 Nov
1983 (stam. fl), J. Brandbyge & A. Barford 42506 (AAU, QCA, QNA, S). Zamora-Chinchipe:
Rd. from Loja to Zamora, km 14, 00° 04'S, 79° 09' W, 2,750-2,770 m, 19-20 Apr 1973
(stam. fl), L. Holm-Nielsen et al. 3965 (AAU, QCA). PERU. Amazonas: Prov. Luya,
Camporredondo-Tullanya, trail to Cerro Huicsocunga, 2,350 m, 3 Sep. 1989 (fl), C. Diaz
& J. Campos 3711 (MO, USM); Parte alta de las Montafias de Galeras, 2,000—2,500 m, 20
Jun 1991 (fl bud), C. Diaz et al. 4448 (BRIT, MO, USM). Ayacucho: Prov. La Mar, E
massif of Cordillera Central, opposing the Cordillera Vilcabamba between Tambo San Miguel,
a and Hacienda Luisiana, 12° 45'S, 73° 53' W, ca. 30 km SW of Hacienda and Rio
Apurimac, 21 Aug 1968 (bisex. ie Dudley 11915 (E, NA, US). Cajamarca: Prov. Cutervo,
10 km NW of Socota, 3,200 m c 1938 (stam. fl.), H. Stork & X. Horton 10134 (F);
San Andrés de Cutervo, Bera er de Cutervo, “Jalca,” trail co Laguna “El Pileo,
2,680 m, 15 Mar 1989 (fl, fr), C. Diaz et al. 3330 (AMAZ, MO, USM); Prov. Jaén, E sie
of Cordillera E of Huancabamba, 2,400-2,600 m, Apr 1942 (fr), A. Weberbauer 6099 (F,
, US). Hudnuco: Prov. Pachitea, region of Pucallpa, W part of Siraé Mountains and
adjacent lowland, ca. 26-28 km ESE from Puerto Inca, 09° 25'S, 74° 43' W, 2,210 m, 15
Aug 1988 (stam. fl), B. Wallnifer 11-10888 (BRIT, MO, W, WU, ae SW slope of Rio
Llulla Pichfs Watershed, on the ascent of Cerros del Sira, top of first cambre between cam
4 (Peligroso) and camp 5 ee 1,680 m, 31 Jul 1969 (ster.), - Dudley 13513 BIS
Lima: Prov. Lima, Lima, without date (fr), J. Pavdn s.n. UX). Madre de Dios: Prov. Manu,
Cerro de Pantiacolla, Rio Plotoa, 10-15 km NNW of Shintuya, 12°35'S, 71°18' W, 1,000—
1,400 m, 15 Dec. 1985 (fr), R. Foster et al. 10860 (F, MO, USM).

GH

Cybianthus pastensis may be easily recognized by its pseudoverticillate leaves

28 SipA 18(1)

with variously serrate or incised margins, the very fine inflorescence rachis
and minute flowers, and the stipitate papillae of the branchlets, leaves, petioles
and inflorescence rachises. The prominent ridges of older branchlets and
the swollen pseudoverticels of leaf scars are also distinctive.

II. Cybianthus subgenus Conomorpha (A. DC.) G. Agostini, Acta Biol.

Venez. 10:150. 1980; Pipoly, Ann. Missouri Bot. Gard. 79:908-957.
eae Conomorpha A. DC., Trans. Linn. Soc. eee ae 17:102. 1834; Conomorpha

ect. Enconomorpha Miq., Src: Surinam. Select. ; 1850: ; Conomorpha subgenus
Enon Mez in Engl., Pllanzenr. IV. 236(Heft 9): i 1902. Type Species: Conomorpha
oblongifolia A. DC. = Cybianthus oblongifolins (A. DC.) G. Agostini (LecroryPE: by

Agostini, - Biol. Venez. 10:151. 1980
Conomorpha sect. Aconomorpha Mig. in Mart., Fl. Bras. 10:304. mete TYPE SPECIES: Conomorpha
heterantha Benth. (LecroryPe: Agostini, Acta Biol. Venez. 10:151. 1980)= Cybianthus
guyanensis (A. DC. ) Mig. subsp. gwyanensis.

Terrestrial dioecious, bisexual, polygamous, or rarely, monoecius shrubs
or trees. Roots positively geotropic. Bark smooth or fissured, brown, or beige,
rarely with significant amounts of cork. Trank distinguishable, leptocaulous,
the growth dynamics following Rauh’s or rarely, Aubréville’s Architectural
Model (Hallé et al. 1978). Branchlets thin to moderately thick, terete or ridged,
densely to moderately covered with ferrugineous stipitate-lepidote scales,
the scales at times appressed, rarely glabrescent. Cataphylls and pseudocataphylls
absent. Leaves alternate, rarely approaching pseudoverticillate (C. peruvianus),
petiolate, covered with ferrugineous stipitate lepidote scales, often glabrescent
above; petioles canaliculate, marginate, or rarely winged. Inflorescence race-
mose, spicate or paniculate, the panicles with racemose branches, rarely a
solitary flower; inflorescence bract small, lanceolate, early caducous; rachis
erect or lax, straight or rarely tortuous, ferrugineous stipitate-lepidote; floral
bracts deltate, lanceolate or ovate, ferrugineous stipitate-lepidote, inserted
at the base of the pedicel; pedicels cylindrical, at times clavate in fruit or
absent, erect, apically recurved, pendent, or nodding, at times accrescent
in fruit. F/owers unisexual, rarely bisexual, dimorphic, (3—)4—5(—6)-merous;
calyx cotyliform, cupuliform, crateriform, urceolate or patelliform, the lobes
valvate, epunctate or with prominent (raised and blisterlike), conspicuous
(readily visible buy flat), or inconspicuous brown, red or black punctations,
the margin entire, rarely crenulate or erose; corolla campanulate to cupuliform,
rarely salverform or tubiform, the lobes erect or reflexed, rarely cucullate,
valvate or imbricate, ferrugineous stipitate-lepidote or glabrous and epunctate
or prominently, conspicuously or inconspicuously pellucid, brown, or black
punctate without, at times with a narrow line of glandular-granules along
the margin, glandular-granulose within, the margins entire or rarely crenulate,
glabrous or rarely glandular-granulose; staminodes resembling stamens but
reduced in size, the tube conspicuous or inconspicuous, adnate to the co-

Pipoty, Cybianthus in Ecuador and Peru 29

rolla, lobate or elobate, the anthers ovate or triangular-ovate, rarely linear-
lanceolate, rarely deltate, usually recurved distally, rarely erect, apically acute,
or apiculate, rarely rounded, the apiculum dorsally, rarely proximally re-
curved or erect, the base cordate, dorsifixed from near base to subversatile,
the connective punctate or not; pistillode conic to lageniform, rarely ab-
sent, translucent-lepidote or glabrous, hollow; pistil obnapiform, rarely conic,
the ovary globose, lobed or with an apical apophysis, the style short, the
stigma capitate-lobate, 2—3-lobed or punctiform; placenta cupuliform or
cotyliform, the ovules 2—4. Fruit subglobose, one(—two)-seede

Cybianthus sabgenus Conomorpha contains 44 species, 10 of which have
been recorded from Ecuador and Peru

KEY TO SPECIES OF CYBIANTHUS SUBGENUS CONOMORPHA

1. Branchlets with erect stipitate aa lepidote scales, their margins
not appressed; leaf blades subbullate to bullate, the secondary veins some-
what to deeply impressed above, eeu raised below; corolla infundibu-
liform or tubiform, or appearing so in
2. Branchlets flexuous, 4-5 mm diam.; leaf blades perpuncticulose above,

sparsely lepidote below, the secondary veins 22—26 pairs, the margin ir-
regular; inflorescence tortuous, pinnately to bipinnately paniculate; co-
rolla lobes prominently keeled, rugose without; anthers ventrally

recurved. 2G: Ee ernies
. Branchlets straight, 2-3 mm diam.; leaf blades not Lo een above
ensely lepidote below, the eae veins 8—19 pairs, the margin feo

ar; inflorescence erect, a simple raceme or poorly ee panicle consist-
ing of basally clustered racemes; corolla lobes flat, smooth or verrruculose
ee Oe anthers dorsally recurved.
3. Secondary veins 12-16; staminal tube epunctate; pedicels cylindric;
” calyx a eae corolla membranaceous, infundibuliform, verruculose
without; fruit smooth, 3. .5 mm diam.; plants of premontane plu-
vial ee subpéramo thickets and upper pluvial cloud forests, 1,000
1,960 m elev

or

; vation 3. C. occigranatensis
eke Scene veins 8—12; staminal tube punctate; eee obconic; calyx
urceolate; corolla carnose, tubiform, smooth without; fruit costate, 7—
15 mm diam.; plants of white sands or on ee 150-180 (—1,500)
m elevation 4. C. spichigeri
les, the margins appressed;
leaf Ste not subbullate or ballate: the secondary veins planar or slightly
raised above, st discernible or slightly raised below; corolla campanu-
pa to cupuliform

Branchlet 1 ferrugineous |

4, Leaf ae coriaceous, rarely chartaceous, the margins subrevolute to
revolute.
ae ie blades densely and prominently pustulate at maturity above, the
secondary veins 24—28 pairs, inconspicuous below; inflorescence spi-
cate (2-)6—16 cm long; flowers subsessile, the pedicels 0.2—0.6 mm
long; eae deeply ae 5. C. lepidotus
5. Leaf b parsely iculate at maturity above,

f

30 Sipa 18(1)

the secondary veins 14—25 pairs, stare! raised below; inflorescence
a raceme or panicle with 2—4 racemes branching from base, (2—)3-—7 (—
8) cm long; flowers pedicellate, Ue pedicels (0.5—)0.9-1.5 mm long;
calyx cotyliform.
6. Branchlets subterete, 2—3 mm diam; leaf blades 1.2—2(—2.5) cm wide,
ree above; petioles S—7(—10) mm long; staminate calyx carnose,
.2-1.8 mm long; stamainace corolla carnose, densely lepidote with-
out, the scales overlapping, 3.2—3.4 mm long, the tube equal to the
staminal tube, the lobes symmetric; anthers ovate; fruit with fleshy
exocarp; plants of montane and cloud forests on sandstone. ..........- 6. C. laetus
6. Branchlets terete, 3-4 mm diam.,; leaf blades (2.6—)3.5—S cm wide,
pusticulate above; petioles 10-15 mm long; staminate calyx
chartaceous, 0.8—1.2 mm long; staminate corolla chartaceous,
glabrous or panel lepidote without, the scales not overlapping,
2.( mm long, the cube shorter than the staminal tube,
the lobes asymmetric; anthers mre es fruit with thin
exocarp; plants of lowland and lower montane forests on white
sands. 7. C. peruvianus
. Leaf blades chartaceous to membranaceous, the margins flac.

SIAR

Leat blades membranaceous to subchartaceous; petioles 5—LO(—12) mm
long; inflorescence a simple raceme or rarely 2-branched at oe 1-3
cm One coe salverform or campanulate; fruit globose.

8. late, 1.5—2 mm diam.; corolla salverform, the stami-
nate 2,.2—2 ran mm; slants of premontane forests on sandstone and
limestone, (244—)400—1,200 m elevation, ..............:0006 8. C. comperuvianus
8. Branchlets terete, 2—3 mm diam.; corolla campanulate, the stami-
nate, 2.8—3.2 mm; plants of lowland igapo forests, 90O—240(—700)
m elevation. . guyanensis subsp. pseudoicacoreus
_Leaf blades chartaceous; petioles (10—)13-17(—22) mm long;
inflorescene a panicle with 2-8 racemes branched from base, 4—8 cm
long; corolla chartaceous, infundibuliform or cotyliform,; fruit de-

~

pressed-globose

9. Branchlets, petioles, abaxial leaf blades, inflorescence and calyx lobes
moderately to densely lepidote, but the not scales overlapping; leaf
blades smooth above at maturity; corolla cotyliform, the lobes ob-
long to oblanceolate, flat, smooch without, conspicuously black

punctate, apically acuminate; staminal and staminodial tube charta-

ceous, conspicuous; anthers and antherodes obcordate, the apiculum
distally recurved; pistillode conic; pistil lageniform .............4 10. C. timanae
9. Branchlets, petioles, abaxial leaf blades, inflorescence and calyx lobes

ey to sean lepidote, the scales overlapping; leaf blades

pusculate above at rity; corolla infundibuliform, the lobes ovate,
verruculose without, eo ite brown punctate, apically
rounded; staminal and staminodial tube membranaceous, inconspicu-
ous; anthers ovate, antherodes subdeltate, the apiculum proximally

—

1. C. cuatrecasasii

inflexed; pistillode lageniform, pistil obnapiform

Ze alate gigantophyllus Pipoly, (Fig. 2A, 10). Candollea 46:41. —
»: PERU. SAN Martin: Pumayacu, between Balsapuerto and Moyobamba, 600—1,200

Pipoty, Cybianthus in Ecuador and Peru Si

Fic. 10. Cybianthus gigantophyllus Pipoly. A. Habit, showing flexuous branchlet, paniculate
inflorescences. B. Pistillate flower, showing urceolate calyx and crenulate corolla lobe mar-
gins. C. Pistillate ower with one corolla lobe removed, showing cucullate corolla lobe
apices, proximally recurved antherodes, and capitate, lobed stigma. A—D, drawn from ho-
lotype, by Peggy Duke. Figure reproduced from Pipoly, 1991.

we
bo

Sipa 18(1)

m, Aug—Sep 1933 ae a G. Klug 3165 (HoLotype: US; isotypes: F, G-2 sheets,
GH-2 sheets, MO, NY.

Tree to 4 m tall. oo ieee prominently ribbed, 4-5 mm diam.,
moderately lepidote. Leaves alternate; blades chartaceous, elliptic, (15.5)17—
27 cm long, (5.9-)7-9.1 cm wide, apically long-acuminate, the acumen
1.2—3.5(-4) cm long, pustulate, perpuncticulose and glabrous above, sparsely
lepidote below, midrib slightly depressed above, prominently raised below,
the secondary veins 22—26 pairs, slightly depressed above, prominently raised
below, the margin irregular, flat, entire; petioles canaliculate, thick, (1.6—)
2.7—4.cm long, ca. 3 mm diam., sparsely lepidote, prominently ridged below.
Staminate inflorescence: unknown. Pistillate inflorescence: a pinnate to bipinnate
panicle, 1.5—2.5 cm long, tortuous, the branches spicate, moderately lepi-
dote; peduncle 0.3—0.5 cm long, floral bracts carnose, deltate, 0.8—0.9 mm
long and wide, apically acute, margin crenulate basally, densely lepidote
above and below; pedicels obsolete. Pssti//ate flowers 4-merous; calyx carnose,
urceolate, 1.6—-1.8 mm long, the tube 0.9—1 mm long, the lobes widely
triangular, 0.5—0.7 mm long, 1—1.2 mm wide, apically acuminate-apicu-
late, the margin regular, entire, lepidote; corolla carnose, campanulate, 2.7—
3.1 mm long, the tube 0.2—0.3 mm long, the lobes erect, 2.5—3 mm long,
apically rounded to obtuse, prominently cucullate, abaxially carinate, apically
rugose and glandular-granulose along the margins without, inconspicuously
black punctate, the margin glandular-granulose, erose-crenulate; staminodes

2.3—2.5 mm long, the staminodial tube membranous, inconspicuous, 0.2—
0.3 mm long, elobate, glabrous, the apical free portions of the filaments
1.2—1.3 mm long, flat, the anthers ovate, 0.8—1 mm long, 0.6—0.8 mm wide,
apiculate, the apiculum ventrally recurved, basally cordate, the connective
epunctate; pistil obturbinate, 1.8—2 mm long, 1—1.3 mm diam., the ovary
1.4—1.6 mm long, the stigma capitate, 3—5-lobed, the placenta cupuliform,
ovules 3, erect, the upper portions exposed. Frvit globose, 4-5 mm long
and in diam., exocarp thin, black, inconspicuously pellucid punctate.

Distribution.—Cybianthus gigantophyllus is known from the headwaters of
the rfos Marafion and Huallaga in San Martin, and the Iquitos area, along
the Rios Napo, Nanay and Amazonas in Loreto, at 130-500 m elevation.

Ecology and conservation status.—Cybianthus gigantophyllus occurs in pri-

mary ferra firme forests, and on white sands (varillal) of lowland Peruvian
Amazonia. Given increasing pressure from deforestation, it should be con-
sidered threatened.

Etymology.—The specific epithet refers to the leaf size, one of the largest
known for the subgenus.

Local names.—Peru: “ukushnum,” “wewé,” “yakisnum,
(Aguaruna).

ye

yaktshnum”

PipoLy, Cybianthus in Ecuador and Peru o15)

Representative specimens examined. PERU. Amazonas: Prov. Bagua, Dtto. Imaza,
ear Aguarana de Putuim (CAMPOU), anexo Yamayakat, Monte Alto de Putuim,
m, 25 Aug 1994 (fl bud), C. Diaz et al. 7007 (BRIT, HUT, MO, USM); Quebrada
Chichi Entsa, Rio Cenepa, 130 m, 7 Jun 1973 (fr), E. Ancuash 580 (AMAZ, M 1);
Vicinity Huampami, 5 km E of Valdivia, 04° 30'S, 73° 30' W, 200-500 m, 12 he ae
(fr), E. Ancuash 1437 (AMAZ, BRIT, MO, NY, US). Loreto: Maquisapa, Upper Rio Nanay,
Jul 1929 (fr), L/. Williams 1182 (F); Prov. Maynas, Dtto. Sta. Marfa de Nanay, Caserfa Mishana,
halfway between Iquitos and Sta. Marfa de Nanay, 03° 50'S, 73° 30' W, 130 m, 25 Feb
1979 (ster.), A. Gentry GJ. Aronson 25044 (AMAZ, MO); Dtto. Las Amazonas, Quebrada
Yanamono, Explornapo Tourist Camp, above mouth of Rio Napo on Rio Amazonas, 9 Nov
se (ster.), A. Gentry et al. 27952 (AMAZ, MO), 25 km NE of Iquitos, along Rio Amazonas,
outhern perimeter pa 110 m, 27 Sep 1990 (ster.), 7. — et al. 12497 (AMAZ, BRIT,
MO, US, USM), 03° 20'S, 72° 55' W, 100-140 m, 15 Feb 1991 (ster.), J. Pipoly et al.
13028 (AMAZ, BRIT, MO, USM), 03° 28'S, 72° 50) = 106 m, 15 May 1989 (ster.), R.
Vasquez et al. 12108 (AMAZ, MO, USM); Exp earn pee Camp, near Sucusari, ae
Rio Napo, 03°20! S, 72°55' W, 100-140 m, 22 Feb 1991 (ster.), J. Pipoly et al. 13284
(AMAZ, BRIT, MO, USM), 23 Feb 1991 (ster.), J. Be et al. 13423 (AMAZ, se MO),
(ster.), J. Pipoly et al. 13426 (AMAZ, BRIT, MO, USM), | Mar 1991 (ster.), J. Pipoly et al.
13931 (AMAZ, BRIT, MO, USM).

Cybianthus gigantophyllus is most closely related to C. occigranatensis (Cuatrec.)
G. Agostini and C. spichigeri Pipoly. However, the large, flat leaves, long
petioles, and tortuous panicles allow for easy recognition. In the original
description (Pipoly 1991), I described the pistillate corolla as tubular, when
it is, in fact, campanulate. When the flower is in bud, the corolla appears
tubular as it longitudinally extends above the calyx, then it gradually opens,
with cucullate apices. Within the tall terra firme forests on lateritic soils, it
may be found above the flood line along small creekbeds.

3. Sera occigranatensis (Cuatrec.) G. Agostini, Acta Biol. Venez.
O215>, 1930; Solas ie occigranatensis Cuatrec., Revista Acad. Colomb. Ci. Exact.
nee 1951. Type: COLOMBIA. Vatie Det Cauca: Cordillera Occidental, W
slope, Rio Digua ee Basin, left bank of Rio San Juan, around Queremil region,
small stream at km 51, 1,540—1,650 m, (stam. fl), J. Cuatrecasas 23734 (HOLOTYPE:

F; isoryPE: COL
Shrub or small tree to 4m tall. Branchlets straight, subterete, 2—3 mm diam.,
densely lepidote. Leaves alternate; blades membranaceous, elliptic to obo-
vate, (4—)7.5—14(—21) cm long, (2.5—)4—5(—7) cm wide, apically acuminate,
the acumen 1.2—1.5(—3.0) cm long, basally acute, decurrent on the petiole,
the midrib impressed above, prominently raised below, the secondary veins
8—12 pairs, deeply impressed above, prominently raised below, the leaf strongly
bullate, adaxial surface smooth, densely lepidote when young, becoming
pusticulate and sparsely lepidote or glabrous with age, abaxial surface densely
lepidote, but the scales not overlapping; petioles canaliculate, 1.0—1.5(—
1.8) cm long, densely lepidote, persistent. Stamnate inflorescence a raceme or
a panicle with 1—3 branches from the base, 4-8 cm long; peduncle, rachis,

34 Sipa 18(1)

branches and pedicels densely lepidote; peduncle 0.1—0.4 mm long; floral
bracts membranceous, ovate, shorter than the pedicels, 0.7—1.1 mm long;
0.4—0.5 mm wide, apically acute, densely lepidote abaxially, the margin
entire; pedicels cylindric, thin, 1.5—6 mm long. Staminate flowers 4-merous;
calyx carnose, cotyliform, 0.8-1.0 mm long, the tube 0.2—0.3 mm long,
the lobes triangular to deltate, 0.5—0.9 mm long, 0.4—0.8 mm wide, apically
attenuate to an acute or round tip, sparsely lepidote without, glabrous within,
conspicuously brown punctate, the margin lepidote; corolla membranaceous,
campanulate, 2.4-2.7 mm long, the tube 0.7—0.8 mm long, the lobes ovate,
1.6—-1.8 mm long, 1.0-1.3 mm wide, apically attenuate to a round tip, verruculose
without, smooth within, sparsely lepidote without toward apex, apically
glandular-granulose within and along margins, conspicuously brown punctate,
the margins entire; stamens 1.8—1.9 mm long, the filaments 2.6—-2.8 mm

long, the tube membranaceous, inconspicuous, adnate to the corolla tube,
elobate, the apically free portions 0.2—-0.3 mm long, the anthers triangular,
0.8—1.0 mm long, 0.5—0.6 mm wide, apically attenuate to an acute , dor-
sally reflexed tip, basally cordate, dorsifixed just above base, the connective
dark, prominently brown punctate; pistillode lageniform, 1.3—1.5 mm long,
densely translucent glandular-lepidote near the base. Bisexual and pistillate
inflorescence: as in staminate but only rarely branched from base, 4—6 cm long;
peduncle 0.1—-0.3 cm long; floral bracts 0.5—0.8 mm long; pedicels 1.5—
3.5 mm long. Bisexual and pistillate flowers as in staminate but calyx 0.8—
1.1 mm long, the cube 0.3—0.6 mm long, the lobes deltate to oblate, 0.4—
0.6 mm long, 0.8-1 mm wide; the margin irregular, entire; corolla as in
staminate but 2.4—2.6 mm long, the tube 1.0—1.1 mm long, the lobes ovate,
1.4—1.6 mm long, 0.8—1.1 mm wide. Bisexual flowers with stamens 1.6—
1.8 mm long, the tube 1.0—1.2 mm long, the apically free portions of fila-
ments (.2—0.3 mm long, the anthers 0.5—0.8 mm long, pistillode 1.5-1.8
mm long. Pzsti/late flowers with staminodes 1.6—1.8 mm long, the tube 1.0—
1.2 mm long, the apically free portion of filaments 0.1—0.2 mm long, the
antherodes (.5—O.7 mm long; pistil obnapiform, 2.4—2.6 mm long, the ovary
1.1-1.2 mm long, 1.1—1.2 mm diam., the style 1.0 mm long, the stigma
capitate, 2-lobed, to 0.2 mm long, the ovules 2—4, buried in the placenta
below apical pores. Frit globose, 2.5—4 mm long, 3.5—4.5 mm diam., the
endocarp smooth, the aril scanty and adnate to both seed and endocarp, the
embryo straight, ca. 3 mm long.

Distribution.—Panama (Darién), Colombia (Cordillera Occidental) and
Ecuador (Esmeralda, Napo, Santiago-Zamora, Sucumbios), at 1 ,QO0-1,960
m elevation.

Ecology and conservation status. —C ybianthus occigranatensis occurs in premontane
pluvial forests, subparamo thickets and in upper pluvial cloud forests. Based
on my observations of populations in subparamo thickets at the Antioquia/

Pipoty, Cybianthus in Ecuador and Peru 2)

Choco interface in the Cordillera Occidental of Colombia, this species tol-
erates disturbance well as long as the soil is not compacted. It is retricted
to areas where rainfall exceeds 5,000 mm annually. At this time, the spe-
cies does not seem to be threatened.

Etymology.—The specific epithet refers to its principal range of distribu-
tion, the Cordillera Occidental of Colombia and adjacent Ecuador.

Specimens examined. PANAMA. Darién: S slope of westermost summit of Cerro Tacaracuna,
massif between Pucro base camp and Tacaracuna summit camp, 1,400-1,600 m, 21 Jul
1976 (stam. fl), A. Gentry et al. 16867 (COL, LL-TEX, MO, PMA). COLOMBIA. Antioquia:
Mcpio Frontino, km 13 Nutibara-La Blanguita Rd., Regién de Murrf, Alto de Cuevas,
06° 44' N, 76° 23' W, 1,990 m, 6 Nov 1988 (fl bud), J. Zarucchi et al. 7201 (BRIT, HUA

; Mpio. Frontino, Vereda Venados, Parque Nacional Las Orquideas, sector Dos Bocas,
confluence of Rio Venados and Rfo Calles, 06° 34' N, 76° 30' W, 29 Oc 1986 (stam. fl), R,
Callejas et al. 2737 (HUA, MO); Mpio. San Lufs, Auitopicts Medellin-Santafé de Bogota,

rf ana, Rd. toward Vereda La Josefina, 18 Dec 1982 (stam. fl), A. Cogollo & C.
Estrada 296 (COL, JAUM, MO); Mpio. Urrao, Parque Nacional Las Orquideas, Vereda
Calles, Permanent Premontane Rainforest Inventory me right bank of Rio Calles, 06°
32'N, 76° 19' W, 1,450 m, 26 Nov 1993 (ster.), J. Pipoly, A. Cogollo et al. 17159 (BRIT,
JAUM, MO), 27 Nov 1993 (ster.), J. Pipoly, A. Ze et a 17182 (BRIT, JAUM, MO),
Range NW of Cabafia de Calles, 1,450 m, 28 Nov 1993 (ster.), A. Cogollo et al. 7529 (BRIT,
JAUM, MO), 1,450-1,500 m, 28 Nov 1993 (ster.), J. Pipoly et al. 17253 (BRIT, JAUM,
MO), (ster.), J. Pipoly et al. 17281 (BRIT, JAUM, MO), 7 Dec 1993 (ster.), J ae et al.
17871 (BRIT, JAUM, MO), 9 Dec 1993 (fl. bud), J. Pépoly et al. 17979 (B T, JAUM
MO), Vereda Calles, Alto de Palmitas, ca. 1 km from Cabafia de Calles, | oa ; 5750 m,
Dec 1993 (ster.), J. Pipoly et al. 17542 (BRIT, JAUM, MO). Chocé: Mpio. Itsmina, een
Raspadura, between Raspadura and Quibd6, split of Rio Atrato and Rio San Juan drainage
basins, ca. 05° 15' N, 76° 38’°W, 18 Apr 1979 (fr), E. Forero & R. Jaramillo 5307 (COL,
MO); Serrania del Darién, along Colombian/Panamanian border, 1,400 m, 20 Jul 1976
(stam. fl, bisex. fl), A. Gentry, H. Leén & L. Forero 16842 (COL, MO); without locality, 1866
(fr), J. Triana 2589 (G). Huila: Rio Suaza, SW of Alejandria, 1,670 m, 23 Aug 1944 (stam.
fl), E. Littl 8532 (COL, US). Quindio: Mariquita, 1866 (stam. fl), J. Triana 2562 (P).
Valle Del Cauca: Cordillera Central, 5 km N of Darién along Rd. toward La Guajira, Upper

» Calima, 03° 58' N, 76° 28' W, 1,550-1,700 m, 24 Jan 1986 (fl bud), B. Stein & L.
McDade 3284 (BRIT, HUA, MO); Finca Zungara, Corregimiento La Divisora, crest of Cordillera
ie W of Cali, 6 km N of Cali-Buenaventura Hwy, 03° 32' N, 76° 35' W, 1,960

, 12 Dec 1985 (ster.), A. Gentry et al. 53167 (COL, MO, US), 24 Mar 1986 (fr), A. Gentry
et o 53551 (COL, MO, US); Rio Digua Drainage Basin, Piedra de Moler, 9O0O-1,180 m,
20 Oct 1943 (pist. fl, fr), J. Cuatrecasas 14918 (COL-2 sheets, F); Rfo Sanquininé, La La-
guna, 1,250—1,400 m, 10 Dec 1943 (stam. fl), J. Cuatrecasas 15658 (COL, F, US); Monte
La Guardia, La Carbonera Range, between Las Brisas and Alban, 1,950—2,000 m, 16 Oct
1946 (stam. fl), J. Cuatrecasas 22131 (COL, FE, US, VEN); San Antonio, W of Cali, 1,900-
2,350 m, 26 Feb 1939 (stam. fl), E. Killip & A. Garcta 33898 (A, BM, COL, F, NY, US); La
Cumbre, 7 May 1922 (stam. fl), F Pennell 5147 (GH, K, NY, US). ECUADOR. Esmeraldas:
San Lorenzo Canton, Reserva Etnica Awd, Parroquia Alto Tambo, Centro de la Unién, Cafion
del Rfo Mira, 00° 52' N, 78° 26' W, 250 m, 22 Mar 1993 (fr), C. G M. Autestia 1313
(BRIT, MO, QCNE). Napo: Carretera Nueva, Cotundo-Coca, 1,130 m, 5 Aug 1984 (pist.
fl, fr), C. Dodson et af. 15115 (MO); Canté6n Archidona, 150 m NE of Caserfo of Huamanj,
right side of Carretera Hollin-Loreto, 00° 43'S, 77° 36' W, 1,200 m, 9 Sep 1988 (fr), F
Hurtado & D. Neill 235 (MO, QCNE), Cordillera de Guacamayos, Rd. to Archidona, Rio

36 Sipa 18(1)
Hollin Pequefio, primary forest on 90° slopes, 00° 38'S, 77° 48' W, 1,900 m, Aug 1990
ah fl), W. Ra is ise as 4899 (BRIT, MO, QCA); Cantén o 3 , Right margin
0 Quijos, : Brava,” of Segundo Pacheco, 00° 12'S, 77° 39' W, 1,800-

oe m, 7- i i. 1990 a W. Palacios 5394 (BRIT, MO, QCNE), — of Volcan Reventador,
— bank of Rio Reventador, between Rd. and trail to crater, 00° 07'S, 77° 36' W, 1,600—

1,850 m, 11 Oct 1990 (seam. fl), - een eres T, MO, QCNE), (eis: fl), W a
6187 (BRIT, MO, QC NE), (fr), Y% — pace MO, QCNE); si any i ete
Coca, Punto oT right margin of Rio Quijos, ca. 10 km $ of Reventador, 00° 1 ale
39' W, 1,500 m, 3—5 Oct 1990 ee fl), W. Palacios S815 (BRIT, MO, QCNE), me 0s! S,
77° 30'W, 14301 m, 6-10 Oct 1990 (pist. f), W. Palacios 6040 (BRIT, MO, QCNE); Yasuni
National Park, Maxus Rd and pipeline construction project, km 15, 00° 31'S, 76° 32' W,
250 m, 30 Jun 1994 (bud) N. Pitman 461 (BRIT, MO). Santiago-Zamora: Between Campanas
and Arenillas, along Rfo Tintas, 10 leagues SE of El Pan, 2,195 m, 13 Jul 1943 (stam. fl),
J. Steyermark 53550 (NY). Sucumbios: Sendero toward Volcan el Reventador from km
100 of Baeza-Lago Agrio Hwy, 1,900 m, 7 Oct 1990 (stam. fl), J. Jaramillo & E. Grijalva
12988 (QCA)

Cybianthus occigranatensis is most closely related to C. timanae Pipoly, but
is easily distinguished by the fewer secondary veins of the coriaceous leaf
blades, the campanulate corolla with verrucose, prominently black punc-
tate lobes and attenuate apices, and obnapiform pistil. The population from
Alto de Cuevas in Antioquia, Colombia, has by far the largest leaves of any
population of this species known thus far. Further study of the population
biologies of Cybianthus montanus ( Lundell) G. Agostini from Panama, C.
occigranatensis, and C. timanae will be necessary to fully resolve the precise
relationships and microecological roles each plays in montane wet and plu-
vial forests.

4, sana spichigeri es Candollea 46:43. 1991. (Fig. 2B, 11).
Type: PERU. Loreto: Prov. Requena, Trocha al Ajuajal, 2 km from Centro Pac
Jenaro Jerrera, right bank of Rio Ucayali, 15 Feb 1982 (stam. fl, fr), R. Spichiger & F.
Encarnacion 1224 (HOLOTYPE: US; isorypes: AMAZ, G, MO).

Tree to 15 m tall. Branchlets thin, straight, terete, 2-3 mm diam., densely
lepidote. Leaves alternate; blades chartaceous, elliptic to narrowly oblanceolate,
(LO) 15-20 em es (3-)5.2—6.5(—7.2) cm wide, apically caudate-acumi-
nate, the acumen 1.9—2.3 cm long, basally acutish to obtuse, not decurrent
on the petiole, bullate, “he midrib and secondary veins strongly impressed
above, prominently raised below, smooth and inconspicuously to promiently
pellucid punctate above, moderately lepidote below, the margin essentially
flat, but very slightly inrolled at the very margin; petioles canaliculate, (1—)
|.2—2 cm long, densely lepidote. Staminate inflorescence: a pyramidal pinnate
pannicle, 1—-4.5 cm long, 1-3 cm wide, peduncle 0.3-1 cm long; branch
bracts chartaceous, linear-subulate, 0.6—1 mm long, 0.1—0.2 mm wide, apically
attenuate, densely lepidote; pedicels cylindrical, (0.8—)1—1.5 mm long. Staminate

flowers 4—S-merous, carnose; calyx suburceolate, 1.3—1.5 mm long, the tube
0.3—-0.5 mm long, the lobes deltate, ca. | mm long and wide, apically acute,

Pipoty, Cybianthus in Ecuador and Peru 37

Fic. 11. neces spichigeri Pipoly. A. Habit, showing minute inflorescences and large,
costate frui Staminate flower in bud, showing suburceolate calyx and tubiform co-
rolla. C. See flower, showing long, prominently lobate staminal tube, cucullate co-
rolla lobes, proximally recurved anthers. A—C, drawn from holotype, by Peggy Duke. Fig-
ure reproduced from Pipoly, 1991.

38 Sipa 18(1)

sparsely lepidote, prominently rugose, with one prominent brown puncta-
tion per lobe, the margins irregular, entire, sparsely lepidote; corolla tubiform,
2.4—2.8 mm long, the tube ca. 0.5 mm long, the lobes oblong, 1.9—2.9
mm long, 0.9—1.1 mm wide, apically acute, prominently cucullate, with
only a few, scattered scales without, glandular-granulose within, the mar-
ginentire, glandular-granulose; stamens 2.2—2.4 mm long, the tube carnose,
conspicuous, 1—1.4 mm long, lobate, the lobes 0.1—0.2 mm long alternat-
ing with the filaments, the apex of the tube and lobes punctate, the apically
free filaments, 0.9—1.3 mm long, the anthers deltate, 0.5—0.7 mm long and
wide, apically apiculate, the apiculum slightly proximally recurved, basally
cordate, the connective dorsally punctate with small brown dots forming a
triangle along connective margin; pistillode conic, 0.5—O.7 mm long, 0.2—
0.3 mm wide, hollow, glabrous, the style conspicuously brown punctate,
the stigma punctiform. Pisti/late iad ie, gamous inflorescence: a raceme, Occa-
sionally a poorly formed panicle of 1-3 racemes branched from base, 0.4—
1.5 cm long, densely lepidote, tardily glabrescent; peduncle 0.1—0.4 mm
long; floral bracts chartaceous, ovate, 0.8—1 mm long, 0.3—0.5 mm wide,
ly attenuate, densely lepidote; pedicels obconic, (0.8—)1.5—2.5(—3) mm
long, to 1.2 mm diam. apically in fruit, densely lepidote. Pisti/late flowers as
in staminate but calyx 1.0—1.2 mm long, the tube 0.4—0.5 mm long, the
lobes 0.6—-0.7 mm long and wide, staminodes and pistil unknown. Frwit

—

apica

depressed-globose, 0.7—0.8 cm long, 0.7-1.5 cm wide, prominently cos-
tate longitudinally, the exocarp costate, inconspicuously pellucid punctate.

Distribution.—Principally known from forests along the rfos Ucayali, Marafion
and Napo Drainage Basin complex, Loreto, Peru, with one disjunct popu-
lation in nearby Morona-Santiago, Ecuador, 150-180 (—1,500) m elevation.

Ecology and conservation status. —Cybianthus spichigert is known from only
a handful of specimens, but is locally quite common. Not enough is known
of the population biology to categorize its conservation status, but its fre-
quency in forest study plots of the Jenaro Herrera Reserve in Peru suggest
it is reproducing and may not be in imminent danger. Cybianthus spichigeri
is a varillal or premontante sandstone species in Peru, and in Ecuador it is
known only from premontane sandstones. Occurrence of this species in the
Cerros del Sira, Peru, reinforces the concept that those mountains contain
many unusual populations of otherwise lowland Amazonian plants.

Etymology.—The epithet commemorates Rudolphe Spichiger, Director
of the Conservatoire et Jardin Botaniques, Chambesy, Geneve, Switzerland.
Dr. Spichiger has devoted much of his career to study of global change,
conservation of biodiversity, and systematics of the genus I/ex. Under his
leadership, the Jardin has maintained active research programs in Paraguay,
Peru, Madagascar, and throughout Europe.

Preoty, Cybianthus in Ecuador and Peru 39

R d. ECUADOR. Morona-Santiago: Cordillera del Condor,
Cuangos, 20 km E of Gualaquiza, near disputed Peru-Ecuador border, 03° 29'S, 78° 14!
1,500 m, 18 Jul 1993 (ster.), A. Gentry 80096 (BRIT, MO, QCNE), 1,470 m, 19 Jul
1993 (infl. bud), A. Gentry 80179 (BRIT, MO, QCNE). PERU. Huanuco: Prov. Pachitea,
region of Pucallpa, W part of “Siraé Mountains,” and adjacent lowland, ca. 24 km SE to 26
km ESE of Puerto Inca, from beginning of rainforest to Campamento Pato Rojo, 09° 27'S,
74° 46' W, 1,380 m, 31 Jan yen (A bud), W. Morawetz & B. Wallnifer es 31188 a
W, WU). Loreto: Prov. May paca IAP Station, 04°10' S, 73°30! W, 150
13 Nov 1984 (fr), R. oe et - 5911 (AMAZ, MO, NY), 6 Jun 1985 (f, R. Vase
al. 6588 ee MO, NY); Rio Nanay, Mishana, 30 km SW of Iquitos, 03° 55'S, 73°
35' W, 150 m, 19 Aug 1978 (fr), R. Foster 4226 (MO, NY, USM), 16 May 1981 (fr), R.
Vasquez a Criollo 1801 (AMAZ, MO, NY), 20 Jan 1985 (fr), R. Vasquez & N. Jaramillo
6137 (AMAZ, MO, NY). Prov. Requena, Reserva Forestal Jenaro Herrera, 04°55'S, 73°45!
W, along Rio Ucayali, 120 m, 1980 (stam. fl), R. Marmillod 9-R-137 (G, US), 1980 (bud),
R. Marmillod 4-R-90, 10 Jun. 1982 (fr), R. Spichiger et al. 1973 (G, US), 24 Feb. 1987
= atry et al. 56531 (AMAZ, MO); Aguajal, 3 km from Centro Forestal Jenaro
Herrera, right margin Rio Ucayali, 18 May 1982 (fr), F Encarnacién 26105 (AMAZ, MO),
22 May 1982 (fr), FE Encarnacién 26200 (AMAZ, MO, NY, US); Arboretum, Centro For-
estal Jenaro Herrera, 180 m, 13 Nov 1981 (fr, stam. fl), R. Spichiger & F. Encarnactén 1027
(AMAZ, G, MO, US)

Cybianthus spichigeri is unique within subgenus Conomorpha because of its
costate fruits. In addition, the subbullate leaf blades, punctate staminal tube,
obconic pedicels, and polygamous inflorescences are also exceedingly rare
characters that allow for easy recognition. Since its description (Pipoly 1991),
an entirely staminate specimen (R. Marmillod 9-R-137) has been located.
While the staminate inflorescence structure is quite different from that of
the polygamous one, flowers of both are identical. The occurrence of this
taxon three times in one inventory conducted at the Jenaro Herrera Re-
serve indicate that the relative frequency of reproductive individuals would
permit a study of the breeding system and population biology for this most
unusual taxon.

a: erie ra lepidotus (Gleason) G. oe Bol. Soc. Venez. Ci. Nat
2:388. 1976. Conomorpha k Gleason, Bull Tertey Bow Gab 18 446: 198
a VENEZUELA. Temmronte FEDERAL vere NAS: Summit of Mt. Duida, 1,500
m, Aug 1928—Mar 1929 (pist. fl), G. H. H. Tate 741 (HoLotyPE: NY, F Neg. 040832;
ISOTYPE: US).

Conomorpha curvivenia Gleason, Bull. Torrey Bot. Club 58:444. 1931. Type: VENEZU-
ELA. TERRITORIO FEDERAL AMAZONAS: Mt. Duida, 1,260 m, Aug 1928—Mar 1929 (stam.

. AH. Tate 927 (HoLotTyPE: NY; tsorype: US).
Gramie lepidota Gleason f. acutata Steyerm., Fieldiana, Bot. 28:465. 1953. Type:
Vv ZUELA. Territorio FeperaL AMAZONAS: Cerro Duida, 2 Dec 1944 (fr), J. Steyermark

58265 (HOLOTYPE: F; ISOTYPE

Shrub or small tree to 6 m tall. Branchlets straight, terete, 2-2.5 mm diam.,
densely lepidote. Leaves alternate; blades elliptic to narrowly elliptic, chartaceous
to coriaceous, (3.1-)5-15 cm long, 1.6—6 cm wide, apically acuminate, the

40 Spa 18(1)

acumen 0).3—2.0 cm long, basally acute to obtuse, midrib depressed above,
prominently raised below, the secondary veins 24—28, inconspicuous above
and below, pustulate and densely lepidote above at first, glabrescent, densely
lepidote below, the scales not overlapping, inconspicuously pellucid punc-
tate, the margin entire, subrevolute to revolute; petioles thin, marginate,
1.0—2.5 cm long, densely lepidote. Staminate inflorescence a spike, rarely two
dubseccile spikes, (2—-)6-16 cm long, 8—15-flowered, peduncle, pedicels, and
axis densely lepidote, the scales not overlapping; peduncle (0.2—)0.5—0.8
mm long; floral bracts ovate to widely ovate, chartaceous, (.5—0.6 mm long,
0.3-0.4 mm wide, apically acute, densely lepidote adaxially, the margin
entire, glabrous. Staminate flowers (4—)5-merous; calyx cupuliform, carnose,
0.9-1.2 mm long, the tube 0.2—0.4 mm long, the lobes deltate to ovate-
triangular, 0.6—-0.8 mm long and wide, apically acute or acuminate, rarely
obtuse, attenuate to a rounded tip, conspicuously brown punctate, the margin
entire, lepidote; corolla cupuliform, carnose, 1.5—2.5 mm long, the tube
0.5—1.0 mm long, the lobes ovate to broadly ovate, 1.2—1.7 mm long, 0.7—
0.9 mm wide, apically attenuate to a round, cucullate tip, at times with a
few, scattered lepidote scales without, glabrous without, glandular-granulose
within over the entire surface, punctations brown, submarginal, the mar-
gin entire, glabrous; stamens 1.5—1.6 mm long, adnate 0.5—1.0 mm to corolla
tube, the staminal tube 0.3—0.4 mm long, carnose, bearing lobes alternat-
ing with the apically free portions of the filaments 0.1—0.2 mm long, the
filaments flat, 0.3—0.4 mm long, erect, glabrous, the anthers dorsifixed less
than 1/4 from base, ovate-triangular, 0.6-0.7 mm long, 0.4—0.5 mm wide,
apically attenuate to a rounded tip, basally cordate, slightly dorsally reflexed,
the connective epunctate; pistillode lageniform, 1.1—1.2 mm long, hollow,
costate basally, sparingly translucent lepidote, pellucid-punctate. Pzstz//ate
inflorescence as in staminate but a spike, (2-)6-16 cm long, 6—10-flowered;
peduncle 0.6-0.8 mm long; floral bracts 0.5—0.6 mm long, 0.3—0.4 mm
wide. Pistillate flowers as in staminate, but staminodes 1.2—1.3 mm long,
adnate 0.5—1.0 mm to corolla tube, the staminodial tube 0.5—0.6 mm long,
carnose, bearing lobes alternating with the apically free portions of the fila-
ments 0.1—0.2 mm long, those apical portions flat, 0.3—0.4 mm long, erect,
glabrous, the actherodes dorsifixed less than 1/4 from base, deltate, 0.7—
0.8 mm long and wide, apically attenuate to a rounded tip, basally cordate,
slightly dorsally reflexed, the connective epunctate; pistil pyriform, 1.4—
1.5 mm long, the ovary 1-1 2 diam., the style not differentiated, the stigma
punctiform, the placenta eartlitoiay eae 2(—3) naked ovules. Fruit globose,
purple at maturity, 0.5—1.0 cm long, 0.6—1.0 cm diam., the endocarp smooth,
the embryo curved, 3.5—4.0 mm long.
Distribution.—Guayana Highland of Venezuela and Brazil, and sandstone

on

Prpoty, Cybianthus in Ecuador and Peru 4]

formations in Bolivia and Peru (reported for the first time here), G(00—2,300
m in Venezuela, 850—950 m in Bolivia, and 760—850 m in Peru.

Ecology and conservation status.—Cybianthus lepidotus is restricted to large
cloud forest formations in transition zones between sandstone and diabasic
intrusions. It is often associated with species of Erythroxylum, which are also
edaphic endemics. It is a widespread, but locally infrequent species and therefore,
should be considered threatened

Etymology.—The epithet refers to the densely lepidote vestiture of the
vegetative and floral parts of the plant.

Specimens examined. PERU. Amazonas: Prov. pee Dtto. Imaza, Comunidad tee
Putuim, anexo Yamayakat, SW of Putuim, 760-850 m, 26 Sep 1994 (stam. fl), C. Diaz et
al, 7252 (BRIT, HUT, MO, USM). BOLIVIA. . Paz: Prov. Nor Yungas, Salley e Rio
Coroico, Sacramento, 10 km NE of Chuspipata on Coroico Rd., 27 Jan 1984 (ster.), A.
Gentry & J. Solomon 44668 (MO), 4 km NE (above) Inabuara, 13.5 km above San Pedro,
1,530—-1,560 m, 22 Jan 1984 (stam. fl), A. Gentry GJ. Solomon 44407 (MO); Prov. Larecaja,
Maipiri, 6 Nov 1926-28 Feb 1927 (stam. fl), O. Buchtien 1758 (HBG, NY, US), eee
10 km S of } pee 850-950 m, 8 Oct-15 Nov 1939 (fr), B. Krukoff 10987 (A, K, MICH
MO, NY, S).

C ee lepidotus, restricted to sizeable cloud forests in transition zones
between sandstone and diabasic intrusions, is most easily recognized by its
long, lax spikes. When sterile, it may be confused with Cybianthus roraimae
(Steyerm.) G. Agost., but may be easily distinguished by the branchlets 2—
2.5 (not 3.5—4.5) mm in diameter. It may also be confused with Cybianthus
punctatus (Mez) G. Agost. and C. cardonae G. Agost. For a discussion of the
differences between these taxa in sterile condition, see Pipoly (1992a).

The area of Bolivia in which this species has been collected is of biogeo-
graphic interest because it also supports several other Guayana Highland
taxa in the Ericaceae and Clusiaceae. Its new discovery in Bagua Province
of Amazonas, Peru, reinforces thematic map data from satellite imagery that
indicated tepuf-like vegetation could be expected in the area. It is interest-
ing that C. Diaz et al. 7252 from Peru, and B. Krukoff 10987 from Bolivia,
are qualitatively and quantitatively identical to specimens of Cybianthus lepidotus
from Cerro Duida, Amazonas, Venezuela.

6. Sea laetus (Mez )G. Agostini (Fig. 2C), Acta Biol. Venez. 10:153.
1980. Conomorpha laeta Mez in Engl., Pflanzenr. IV. 236(Heft 9):259. 1902. Type:
PERU. Amazonas: Taulia, without elevation or date (stam. fl.), A. Matthews 1561
(HOLOTYPE: K; ISOTYPE: K
Shrub or small tree to 2 m tall. Branchlets straight, subterete, 2—3 mm diam.,
densely lepidote. Leaves alternate; blades coriaceous, obovate, 3—8(—12) cm
long, 1.2—2(-5.0) cm wide, apically acute or short-acuminate, basally cu-
neate, decurrent on the petiole, midrib depressed above, prominently raised
below, the secondary veins 14-18 pairs, planar above, conspicuous below,

4? Sipa 18(1)

glabrous and smooth above at maturity, densely lepidote below, the mar-
gin revolute; petioles marginate, 0.5—1 cm long, densely lepidote. Stamz-
nate inflorescence: a raceme or panicle with 2 racemose branches from base,
2—3 cm long; rachis, and pedicels densely lepidote; peduncle 0.1—0.3 cm
long; pedicels cylindrical, 0.9-1.5 mm long; floral bracts chartaceous, nar-
rowly ovate, 1-2 mm long, densely lepidote adaxially. Staminate flowers 4-
merous; calyx carnose, cotyliform, 1.2—1.8 mm long, sparsely to densely
lepidote without, glabrous within, the tube 0.2—0.3 mm long, the lobes
triangular to deltate, 0.8—1 mm long, 0.6—-0.8 mm wide, apically attenu-
ate to a rounded tip, inconspicuously pellucid punctate, the margin entire,
lepidote; corolla carnose, campanulate, 3.2—3.4 mm long, the tube 1.8—2
mm long, densely lepidote without, the scales overlapping, glabrous within,
the lobes ovate or ovate-triangular, 0.8—0.9 mm long, 0.3-0.4 mm wide,
symmetric, apically attenuate to a rounded tip, inconspicuosly pellucid punctate,
densely lepidote without, glandular-granulose within, the margin entire,
glandular-granulose; stamens 2.6—2.7 mm long, the tube carnose, conspicuous,
equalling the corolla tube, 1.8—2 mm long, lobate, the lobes alternating
with the filaments ca. 0.1 mm long, the apically free portions of the fila-
ments flat, 0.1—O0.2 mm long, the anthers dorsifixed, ovate, 0.6-0.9 mm
long, |.8—1.9 mm wide, somewhat curved distally, apically apiculate to
subapiculate, basally cordate, the connective inconspicuously brown punc-
tate; pistillode conic, 1.5—2 mm long, translucent glandular-lepidote ba-
sally, hollow. Pist//late inflorescence: as in staminate but 3—3.5 cm long; pe-
duncle 0.1—0.4 cm long; pedicels cylindrical, 0.9-1.5 mm long; floral bracts
chartaceous, linear-lanceolate, 1.3—1.5 mm long, densely lepidote adaxially.
Pistillate flowers as in staminate but calyx 1.2—1.4 mm long, sparsely to densely
lepidote without, glabrous within, the tube ca. 0.2 mm long, the lobes deltate,
1—1.2 mm long and wide, apically attenuate to a rounded tip, inconspicu-
ously pellucid punctate, the margin entire, lepidote; corolla, staminodes
and pistil unknown. Frvit globose, S—8 mm long and in diam., exocarp carnose,
ack.

Distribution.—F*astern slopes of the Andes, Colombia, Peru and Bolivia,
1,980—2,850 m.

Ecology and conservation status. —The species occurs in primary cloud for-
est, a life zone being cleared rapidly for cultivation throughout the Andes,
which may account for the paucity of collections. Within subgenus Conomorpha,
aaah laetus is the species most in oan of extinction,

Ety ight,” and probably
refers to che plant’s aesthetically ane appearance, having the same general
form as many species of Vaccinium, Myrsine dependens, other Ericaceae, and
other diminutive Andean shrubs. The thick juicy exocarp is said to be very
tasty although slightly acidic (T. Dudley, pers. comm.).

Pipoty, Cybianthus in Ecuador and Peru 43

Representative specimens examined. COLOMBIA. Boyaca: Arcabuco, NE of town, 2,650
m, 11 Nov 1965 (stam. fl), L. Uribe s.n.(COL); Sierra Nevada del Cocuy, path from Laguna
to Cobug6n, near Alto del Oso, 2,900 m, 27 Aug 1958 (stam. fl), BP. Grubb et al. 744 (K).
PERU. Amazonas: Prov. Luya, Dtto. Camporredondo, Anexo Tullanya, between Pajaco
Tigre and Palma, 06° 04' 35"S, 78° 21' 45" W, 2,500—2,600 m, 9 Dec 1996 (fr), J. Campos
et al. 3161 (BRIT, HUT, MO, USM); Along Rd. E of Chachapoyas between Pipos and
Molinopampa, 06°15'S, 77°40! W, 1,980—2,340 m, 14 Feb 1985 (pist. fl, fr), J. Luteyn &
E. Cotton 11414 (NY, TEX, US, USM); E of Chachapoyas, 2,000 m, without date (stam.
fl), A. Weberbauer 4354 (G). Cusco: Prov. La Convenci6én, Cordillera Vilcabamba, 12° 37'
S, 73° 32' W, ceja and cumbre, 2,550 m, 3 Jul 1968 (pist. bud), 7. Dudley 10690 (F, NA,
USM), 5 Jul 1968 (pist Here). E ee 10803 (&, NA, USM). Huanuco: Prov. Hudnuco,
Carpish Hills, trail to summit from W entrance, 2,700—2,850 m, 09° 42'S, 76° 05' W, 2

Mar 1985 (stam. f1.), B. Stein & C. eo 292 (MO, USM).

Cybianthus laetus is easily recognized by its small leaves, short petioles,
subsessile anthers and thick, juicy exocarp. It is known outside of Peru from
only two other collections, one from the department of Boyaca, Colombia
and the other from Nor Yungas, Bolivia. A recent collection from Luya Province
in western Peru (J. Campos et al. 3161) is referred here because of its thin,
angulate branchlets, and the thick exocarp of its fruits, despite the larger,
thinner leaves. It is expected in Ecuador, in either Napo or Santiago-Zamora
Provinces.

7. Cybianthus peruvianus (A. DC. ) oe ig. oe in Mart., Fl. Bras.
10:298. 1856. eae beruviana A. n. Sci. Nat., Bot. ser 2, 16:92.
1841. Peckia peruviana (A. DC. ) Kuntze, ae ee Pi His Hor Type: PERU.
AMAzONAS: Moyobamba, pe a am. fl), A. Mathews s.n, (HOLOTYPE: G-DC; IsoTyPEs:

,G sH, K)

Conomorpha weberbaueri Mez, Repert. Spec. Nov. Regni Veg. 3:101. 1906. Type: PER
Loreto: In mountains near Moyobamba, 1,300 m, 28 Aug 1904 (stam. fl), A. ‘eae
4668 (HOLOTYPE: B-destroyed; Lecrorypr, here designated: F).

Shrub or small tree to 16 m tall. Branchlets straight, terete, 3-5 mm diam.,
densely lepidote. Leaves alternate, at times approaching eae
at some nodes; blades coriaceous, elliptic to obovate, 7—-12.7 cm long, (2.6—)
3.5-5 cm wide, apically acuminate, the acumen 1.2—1.6 cm long, basally
acute to obtuse, decurrent on the petiole, midrib depressed above, promi-
nently raised below, the secondary veins 10—25 pairs, prominently raised
below, densely lepidote at first, then pusticulate above at maturity, sparsely
lepidote below, the margin revolute; petioles canaliculate, 1-1.5 cm long,
densely lepidote. Staminate inflorescence: a panicle with 1—4 racemes branch-
ing from the base, (2—)3—8 cm long; peduncle, rachis and pedicels densely
lepidote; peduncle 0.1—0.2 cm long; floral bracts chartaceous, ovate, 0).7—1
mm long, 0.5—0.6 mm wide, apically attenuate, densely lepdote abaxially;
pedicels cylindrical, 0.5-1(-1.2) mm long. Staminate flowers 4-5-merous;
calyx green, cotyliform, 0.8—1.2 mm long, lepidote without, glabrous within,
the tube 0.3—-0.4 mm long, the lobes ovate to deltate, 0.7—0.8 mm long,

44 Stipa 18(1)

0.5—0.8 mm wide, apically attenuate to a rounded tip, the margin lepi-
dote, inconspicuously brown punctate; corolla translucent green, chartaceous,
campanulate, 2—2.6 mm long, the tube 0.6—-1.2 mm long, glabrous or sparsely
lepidote externally, glabrous internally, the lobes ovate to narrowly ovate,
1.2—1.8 mm long, 0.5—1.1 mm wide, asymmetric, apically rounded or at-
tenuate to a rounded tip, sparsely lepidote externally, the scales not over-
lapping, sparsely glandular-granulose internally, inconspicuously brown punctate,
the margin glandular-granulose; stamens 1.4—2 mm long, the tube chartaceous,
conspicuous, longer than the corolla tube, 0.8—1.2 mm long, elobate, the
apically free portions of the filaments 0.2—0.4 mm long, the anthers linear-
lanceolate, 0.7—0.9 mm long, 0.2—0.3 mm wide, somewhat recurved dis-
tally, attenuate toa round or acute tip, the connective conspicuously brown
punctate; pistillode obclaviform, 1—-1.5 mm long, translucent glandular-
lepidote basally. Pzstz//ate inflorescence: like the staminate but, 2-7 cm long;
peduncle 0.1—0.2 cm long; floral bracts 0.7—1 mm long, 0.4—0.5 mm wide;
pedicels 0.2-0.5 mm long. Pastil/late flowers as in staminate but calyx 0.9—
1.1 mm long, the tube 0.3—0.7 mm long, the lobes deltate, 0.5—0.6 mm
long, 0.4-0.5 mm wide, apically subacute to obtuse; corolla 1.3—1.7 mm
long, the tube 0.2—0.3 mm long, the lobes ovate, 1—1.4 mm long, 0.7—0.9
mm wide, apically obtuse; staminodes 1—1.2 mm long, the tube longer than
the corolla tube, 0.3—0.4 mm long, the apically free portions of the fila-
ments 0.2—0.3 mm long, the antherodes 0.6—0.7 mm long, 0.2—0.3 mm
wide, pistil obnapiform, 1.3—1.4 mm long, 1—1.1 mm diam., the ovary 0.6—
0.7 mm long, translucent glandular-lepidote basally, the style 0.5—0.6 mm
long, conspicuously brown punctate, the stigma punctiform, the placenta
globose, ovules 3, apically exposed. Frait subglobose, 0.3—0.8 mm long,
0.4-0.9 mm diam., the exocarp thin, black.

Distribution.—Amazonian Ecuador through Peru to Bolivia, at 122—1,500
m elevation.

Ecology and conservation status. —Cybianthus peruvianus occurs on the east-
ern slopes of the Andes in moist or wet lowland and premontane forests on
white sands, especially in transition zones, where brownish sand-clay mix-
tures occur.

Etymology.—The epithet refers to the type locality, in (Moyobamba) Peru.

Local name.—Petu: “Tarrafa caspi.” (Quichua), “uchi yacushnum” (Aguaruna).

1. ECUADOR. Napo: Cantén Aguarico, Reserva Faunistica

Cc gabe no, Laguna Zancudo Cocha ey SE side of Laguna, 00° 33'S, 75° 32' W, 22
m, 28 Sep 1991 (fr), W. Palacios et al. 7761 (BRIT, MO, QC abe Cann eet Sector
teem 20 km N of Coca, PAL oe property, 00° 20'S, > O5' W, 250 m, 3—
v 1989 (fr), E. Gaidifio 137 (BRIT, MO, QCNE); ies to ae ope. 28 Apr
ee fl), J. Jaramillo 8522 (QCA). Zamora-Chinchipe: Cant6n Nangaritza fo
Miazi, along Rio Nangaritza, 900 m, 19 Feb 1994 (fr), H. van ne Werffet al. 13280 (BRIT
MO, QCNE); Hill above military post, 04° 18'S, 78° 40' W, 1,000 m, D. — GW.

=~
-

Prpoty, Cybianthus in Ecuador and Peru 45

Palacios 9615 (BRIT, MO, QCNE), 04° 16'S, 78° 42' W, 970 m, 20 Oct 1991 (fr), W.
Palacios et al. 8486 (BRIT, COL, MO, QCNE). PERU. Amazonas: Along Rio Marafion,
near confluence with Rio Santiago, 1924 (stam. fl.), G. Tessmann 3525 (B, G); Prov. Bagua,
Dtto. Imaza, NE region of Rio Marafion Basin, Comunidad Kampaenza, along
Quebrada Shimutaz, Rfo Marafion, 04° 55'S, 78° 19' W, 320 m, 9 Sep 1994 (pist. fl, fr),
N. Jaramillo et al. 436 (AMAZ, BRIT, HUT, ae USM), 09 Oct 1995 (fr), N. Jaramillo G
D. Chamik 813 (AMAZ, BRIT, MO, USM); Comunidad Aguaruna de Putuim (CAMPOU),
anexo Yamayakat, Monte Alto de Putuim, 450 m, 25 Aug 1994 (fr), C. Diaz et al. 6993
(BRIT, HUT, MO, USM), 660-760 m, 21 Sep 1994 (stam. fl), C. Diaz et al. 7170 (BRIT,
MO, USM); Cerros de Putuim, 05° 03' 20"S, 78° 20! 23" W, 350 m, 13 Jun 1996 (fr), R.
ae et al, 21131. Huanuco: Prov. Pachitea, region of Pucallpa, W part of Siraé Moun-
ns and adjacent lowland, 20-24 km SE of Puerto Inca, Campamento Oro, 09° 29'S, 74°
50 me to Campamento Sir4, 800 m, 17 Jul 1988 (stam. fl), W. Morawetz G B. Peale

2-19188 (BRIT, W, WU). Loreto: Prov. Loreto, Nauta, 04° 32'S, 73° 35' W, 160 m
ae 1984 (fr), R. Vasquez GN. Jaramillo 5060 (AMAZ, MO, USM); Zuingaru Cocha, iF
km SE of Iquitos, 9 Sep 1964 (stam. fl), C. Dodson 2821 (AMAZ, MO, US, USM); Prov.
Maynas, Moropon, lower Rio Nanay above Bellavista, 29 Aug 1968 (fr), $. McDaniel 10942
(AMAZ, IEB, MO); aie ne he rada Blanco Biol. Station, ween II Deis Blanco,
Tahuayo River, 04° 23'S, 73° 1 4 Apr 1985 (fr), J. Castro 27 (A Z, MO, US, USM);
Dtto. Iquitos, Hwy to Sto. a S, a in front of “Chaparal” ee farm, 140 m, 16
Dec 1983 (stam. fl), a Rimachi ie (AMAZ, IBE, MO, US); Estacién Experimental HAP
Allpahuayo, 21 km S of Iquitos,04° 10'S, 73° 30' W, 160 m, 18 Sep 1990 (ster.), J. Pipoly
et al. 12112 (AMAZ, MO, US, USM), 220 m, 19 Sep 1990 (stam. fl), J. Pipoly et al. 12210
(AMAZ, MO, NY, USM); Deto. Sta. Maria de aie LO km W of Caserfo Mishana, in
Cocha Yarama Reserve, 03° 55'S, 73° 35' W, 130 m, 14 Mar 1991 (ster.), J. Pipoly et al.
14994 (AMAZ, MO, US, Ve) (ster.), J. Pipoly val ee MO, USM); Mishana,
along Rio ct O32 1S. 73732, 130 my. 22 1986 (stam. bud), R. Vasquez et al.
7503 (AMAZ, MO, ae USM), 8 Sep 1990 (fr), R. Moe et al, 14335 (AMAZ, BISH, F,

, NY, TEX, US, USM); Mishana,0 3° 52! S, 73° 30' W, 140 m, 4 Jan 1983 (fr), A.
Can et al. 39040 ee MO, USM); Dtto. bea eas Almendras, Arboretum Ciencias
de Ingenierfa Forestal, UNAP, 03° 48'S, 73° 25' W, 122m, 4 Sep 1992 (ster.), - Grades
et al. 4711 (AMAZ, BRIT, MO, USM), Puerto Almendras, along Rio Nanay, 03° 45'S

3° 25' W, 122 m, 30 Oct 1984 (fr), R. Vasquez & N. Jaramillo 5867 ae ee NY,

M), 29 May 1986 (fr), R. Vézquez G N. Jaramillo 7570 (AMAZ, on USM), 30
a 1986 (stam. bud), R. Vasquez & N. Jaramillo 7587 (AMAZ, MO, US, US oo.
IMARPE tract, along Rd. 13 km from Iquitos, 14 Jul 1976 (fr), E Encarnacion - (AMAZ,
US); Prov. Requena, Dtto. Sapuena, Jenaro Herrera, Rio Ucayali, 0 04° 55'S, 73° 40' W,
160 m, 16 Aug 1994 (stam. fl), R. Ort/z et al. 98 (AMAZ, BRIT, MO, USM). Madre de
Dios: Prov. Tambopata, Santuario Nacional Pampas del Heath, Quebrada Palma Real Grande,
12° 57'11"S, 68° 54' 48" W, 210 m, 21 Apr 1996 (fr), M. Sas & D. Castro 623 (BRIT,
MO, USM), Puesto Enahuipa, 12° 39' 23"S, 68° 44' 13" W, 210 m, 25 Apr 1996 (fr), M.
Aguilar & D. Castro 655 (BRIT, MO, USM). San Martin: Le oe Martin, trail to telev
sion antenna, km 17.5 of Tarapoto-Yurimaguas Rd., 2.5 km N of Cataratas de reer ned
06° 27' S, 76° 21' W, 850-1,200 m, 7 Sep 1986 (pist. fl, fr), S. Knapp 8290 (MO, US,
USM).

Cybianthus peruvianus is most closely related to C. comperuvianus Pipoly
(herein described), but may be recognized by its longer, canaliculate peti-
oles, revolute leaf margins, shorter pedicels, asymmetric, lepidote corolla
lobes, and linear-lanceolate anthers. Field studies near Iquitos have shown

46 Sipa L8(1)

Fic. 12 ees comperuvianus Pipoly. A. Habit, showing inflorescence of racemes or malformed
panicles. B. Staminate flower and axillant bract, showing floral densely lepidote floral bract
as long as pedicel, and lepidote calyx margins. C. Opened staminate flower, showing staminal

tube longer than apical free portions of the filaments, lobes glandular-granulose adaxially
and crenulate margins. A—C, drawn from holotype, by Linny Heagy.

Prpoty, Cybianthus in Ecuador and Peru 47

it is a ridgetop species in the lowlands, with a density of approximately 2—
4 individuals per hectare.

8. “oa comperuvianus Pipoly, sp. nov. (Fig. 2E, 12). Type: BRA-
Maro Grosso: Sta. a da Chapada, 1903 (stam. fl), G. re we (HOLO-
TYPE: S; ISOTYPES: G, GH, R, S).

Cybianthus comperuvianus eae ex Pipoly in Killeen et al., Guia Arb. Boliv. 570. 1993.
nom. nud. Agostini (1972) provided the first description of this species in his disserta-
tion, but never published it. I subsequently annotated herbarium specimens with the
name, fully intending to publish it, but it was still not validly published when it
appeared in Guia de Arboles de Bolivia, without Latin description or reference to type.
Presumably, the name was obtained by them based on herbarium determinations, and
thus a zomen nudum and invalid. Validation of the name is effected here,with the cita-
tion of holotype above and the Latin diagnosis, and accompanying description, pro-

vided below.

Species haec cum C. p nfusus est, sed laminis membranaceis (non

eee secus Tortie. eplanis tae revolutis), ie marginaee (non canaliculatis) 0.5—
1 (nec 1—1.5) longis, pedicellis 1.5—2.5 (non 0.7—1.0) mm longis, corollis staminaribus
ee (non campanulatis) statim diagnoscenda.

Shrub or small tree to 6 m tall. Branchlets angulate, 1.5—2 mm diam., densely
lepidote. Leaves alternate; membranaceous, elliptic, narrowly elliptic, to obovate,
9-13(-17) cm long, 3—5(—6) cm wide, apically acuminate, basally acute,
decurrent on the petiole, the midrib slightly depressed above, prominently
raised below, pusticulate above, densely lepidote below, the secondary veins
20-30 pairs, the margin flat; petioles marginate, 0.5—1 cm long, densely
lepidote. Staminate inflorescence: a raceme, rarely a malformed panicle with 2
branches from the base, 1—3 cm long; peduncle 0.1—0.5 cm long; floral bracts
membranaceous, ovate, 1.3—1.5(—2.0) mm long, ca. 0.6 mm wide, subglabrous,
sparsely lepidote above and densely lepidote below, apically acute, slightly
shorter than or as long as the pedicels, entire; pedicels cylindrical, 1.2—2.5
mm long, sparsely lepidote. Staminate flowers whitish-green, 4(—5)-merous,
membranaceous; calyx cupuliform, 1—-1.2 mm long, the tube 0.3—0.6 mm
long, the lobes triangular to deltate, 0.6—1 mm long, 0.5—0.7 mm wide,
apically attenuate to a rounded tip, sparsely lepidote without, glabrous within,
conspicuously brown punctate, the margin entire, sparsely ferrugineous-
lepidote; corolla salverform, 2.2—2.6 mm long, the tube 1-1.2 mm long,
the lobes ovate to ovate-triangular, 1.2-1.6 mm long, 0.8—1 mm wide, apically
rounded or attenuate to a rounded tip, glabrous without, glandular-granulose
throughout within, the margin minutely crenulate, glandular-granulose,
conspicuously brown punctate; stamens 1.8—2.2 mm long, adnate 1.2—1.5
mm to corolla tube, staminal tube 1.6—-1.9 mm long, elobate, longer than
the apically free portions of the filaments 0.2—-0.3 mm long, the anthers
deltate to triangular, slightly distally curved, 0.7-0.8 mm long, 0.5—0.6
mm wide, apically apiculate, basally broadly cordate, the connective incon-

48 Stipa 18(1)

spicuously brown punctate; pistillode lageniform, 1.0-1.2 mm long, 0.3—
0.4 mm wide, the ovary 0.4—0.5 mm long, densely translucent glandular-
lepidote near base, the style 0.5—0.6 mm long, the stigma punctiform. Pis-
tillate inflorescence as in staminate but pedicel 1.5—2.5 mm long. Pistillate
flowers as in staminate but calyx 1.0-1.2 mm long, the tube 0.3—0.6 mm
long, the lobes deltate, 0.6—1 mm long and wide, corolla, staminodes and
pistil unknown. Fruit globose, 5-7 mm long, 6-8 mm diam.

Distribution.—Ecuador southward to Bolivia and adjacent Brazil, 400—
1,200 m elevation.

Ecology and conservation status. —Cybianthus comperuvianus occurs in primary
premontane forests on sandstone and limestone, in relatively sparse popu-
lations (teste collectore). Therefore, it should be considered threatened.

Etymology.
has long been confused with C. perwvianus.

Local name.—Peru: “wewe” (Jivaro); “uchi apikna” (Huambisa); “Cumalilla”
(Spanish).

Paratypes. ECUADOR. Morona-Santiago: Sitio La Planada, 01° 46! $, 77° 57! W,
900 m, 21 Sep 1993 (stam. fl), W. Palacios 11407 (BRIT, MO, QCNE). PERU. Amazonas:
Near Yucui Entsa, 6 hrs. from Camino de Kusu, 300 m, 11 Mar 1973 (fr), R. Kayap 558
(MO, NY, USM), (fr), E. Anciash 93 (AMAZ, BRIT, EF, LL-TEX, MO, NY, US, USM); Quebrada
Huampami, Tseasim, near Nayumpim, 244 m, 3 Apr 1973 (fr), E. Ancuash 141 (AMAZ,
F, LL-TEX, MO, US, USM); Quebrada Yuctui Entsa, 305 m, 12 Apr 1973 (fr), E. Ancuash
220 (AMAZ, F, LL-TEX, MO, US); Prov. Bagua, Dtto. Imaza, NE Region of Rfo Marafion
Drainage Basin, Comunidad de Yamayakat, Rio Marafion, 04° 55'S, 78° 19' W, 320 m, 8
Aug 1994 (stam. fl), N. pes et al. 321 (BRIT, HUT, MO, USM). Ayacucho: Tambillo
ee Boa 27 Jul. 1878 ,C. Jelski 3600 (PR, W). Hudnuco: E of Tingo Maria, 5

. L972 (stam. f1.), T. pao a 7 (F, USM, MO); Agua Blanca, trail to Monz6n, 9 Feb
on ee fl.), J. Schunke 1049 (AMES, MO, NY, S, USM, VEN); vicinity Rondos, 24
Mar. 1962 (stam. f.), J. Schunke 5881 (F, US, USM); oo Pachitea, region of Pucallpa, W
part of Sira Mountains and ee lowland, 20-24 km SE of Puerto Inca, Campamento
Oro, 09° 29'S, 74° 50! W, to | Sira, 800 m, 17 Jul 1988 on fl), B. Wallnifer
12-17788 (BRIT, MO, W, Se 30) Aug 1988 (fr), W oo . & B. Wallnifer 13-30888
(BRIT, W, WU), from Campamento Sird, 09° 28'S, 74° W, SE to valley of Rio Negro,
650 m, 11 Aug 1988 (fr), W. Morawetz & B. Wallnifer Pe 11888 (BRIT, MO, W, WU).
Junin: E of Quimirf Bridge, near La Merced, 800—1,300 m, | Mar 1929 (stam. fl.), E.
Killip & A. C. Smith 24011 (F, NY, US); La Merced, Hacienda Schunke, 27 Aug—l Sep
nee (stam. fl.), J. Machride 5677 (F); Pichis Trail, San Nicolas, 1,100 m, 4 Jul 1929 (stam.
E. Killip & A. C. Smith 26073 (F, NY, US), Sta. Rosa, 625-900 m, 6 Jul 1929 (stam.
- eee it. F, NY, US S, USM): Puerto Yessup, a m, 10 Jul 1929 (stam. f1.), E. Killip
GA. C. Smith 26286 (NY, US); Puerto Bermudez, 375 m, 14 Jul 1929 (stam. fl.), E. Ké/lip
& A.C. Smith 26464 (NY, US), 26 ey (NY, US), 26563 (NY, US). Pasco: Prov. Oxapampa,

Pichis Valley, San Matias Ridge, 10-12 km SW of Puerto Bermudez, above Sta. Rosa de
Chiris, aa to Loma Linda, 10° . . 75° 00' W, 500 m, 29 Sep 1982 (fr), R. Foster et al.
8962 (FE, MO, USM). Puno: Below San Gabén on Rio San Gabon, 300—1,000 m, 17—24

Jul 1978 ee bud), Mf. Di/lon et al, 1219 (BRIT, E MO, USM). San Martin: Prov. Mariscal
Caceres, Dtto. Tocache Nuevo, Palo Blanco near Fundo de Manuel Aranjo, 700-800 m, 1

Prpoty, Cybianthus in Ecuador and Peru 49

Mar oe ee Schunke pee (AMAZ, BRIT, F, MO, NY, US); without specific locality,
1778-1788 (stam. fl.), Ruéz L. & J. Pavin 5/36 (F, MA). BOLIVIA. La Paz: Mapiri Re-
gion, 1926 ce 20; Buchvien 1753 (F, GH, HBG, NY, US); Tuiri, near Mapiri, 490—
750 m, Sep 1939 (stam. fl.), B. Krukoff 10930 (A, G, GH, MO, MICH, NY, S, U, UC, US).
Santa Cruz: Velasco; Parque Nacional Noel Kempff Mercado, Campamento las Gamas,
14° 48' 41" S, 60° 23' 45" W, 850 m, 26 Mar 1993 (fr), L. Arroyo & K. Keill 164 (BRIT,
MO, USZ); Campamento Huanchaca, 13° 54'S, 60° 48' W, 650 m, 17 May 1994 (stam.
fl), L. Arroyo et al. 674 (BRIT, MO, USZ). BRAZIL. Mato Grosso: Mpio. Cuiaba, Burity,
NE of Cuiaba, 750 m, Jul 1927 (stam. fl.), B. Collenette 113 (NY); Chapada dos Guimraes,
Cachoeira Véu de Noiva, do Rio Coxipozinho, 15° 30'S, 55° 45' W, 21 Oct. 1985 (fr), J.
Pirani 1326 (NPA, MG, NY, SP), 720 m, 16 Oct 1973 (fr), G. Prance et al, 19075 (AAU,
5 , K, MG, NY, K, S, SP, U, US); Sta. Anna da Chapada, 1902 (stam. fl), G. Ma/me
2048 (S, UPS), 1903 (fr), G. ae 3483 (G, GH, R, S), 1827 (stam. f1.), L. Riedel 959
(LE, NY, US), 1902 (fr), A. t 322 (BM). Rond6nia: 1 km NE of Ariquemes, Porto
Vehlo-Cuiaba Hwy., 13 Aug eee (fr), E. Forero & L. Wrigley 7035 (MG, MO, NY).
Cybianthus comperuvianus was first recognized as a novelty by Agostini (1972).
Despite the fact that over 25 years has past since its first recognition, pistil-
late flowers are still unknown, reinforcing the concept that within the ge-
nus they are ephemeral (Pipoly 1983a, 1992). Cybianthus comperuvianus is
most closely related to C. perwvianus but is easily distinguished by its long
pedicels, salverform staminate corolla, and membranaceous leaves.

9. Cybianthus let aae (A. DC.) Mig. in Mart. subsp. pseudoicacoreus
(M Mar t.) Pipoly, comb. et stat. nov. (Fig. 2F). ee

Mig. in Mart., Fl. Bras. 10:284. 1856. He sii (Miq. in Mart.) M
in eae Baenceor IV. 236(Heft 9):261. . Cyb Aeon (Mig. in
rt.) G. Agostini, Acta Biol. Venez. eee 1980. TYPE: BRAZIL. AMAzonas:
ee near Rio Japurd, Jan 1820 (pist. fl, fr), C. Martius s.n. on here desig-

nated: M; ISOLECTOTYPE: M

Shrub or tree to 7 m tall. Branchlets terete, 2-3 mm diam., densley lepi-
dote. Leaves alternate; blades membranaceous to chartaceous, narrowly obovate
to elliptic, symmetric, (8—)9—12(-18.5) cm long, 3-4.5(-5.7) cm wide, apically
abruptly acuminate to caudate, the acumen 1—2 cm long, basally cuneate,
decurrent on the petiole, midrib flat or slightly depressed on the upper surface,
prominently raised below, the secondary veins 12—25 pairs, pustulate above,
sparsely lepidote below, the margin flat; petioles canaliculate, 0.5—0.8(—-1.5)
cm long, densely lepidote. Staminate inflorescence: a raceme or panicle with
1—2 branches from base, 1-3 cm long; peduncle, rachis and pedicels densely
lepidote; peduncle 0.1—0.2 mm long; floral bracts chartaceous, narrowly
ovate, longer than the pedicels, 1.3—1.5 mm long, 0.2—-0.3 mm wide, apically
attenuate, densely lepidote abaxially; pedicels cylindrical, 1—1.2 mm long.
Staminate flowers 4-merous; calyx carnose, subcupuliform, 1—1.2 mm long,
sparsely lepidote without, glabrous within, the tube 0.3—0.4 mm long, the
lobes triangular, 0.8—1(—1.2) mm long, 0.5—0.7 mm wide, apically attenu-
ate to a rounded tip, conspicuously brown punctate, the margin lepidote;

SO Sipa 18(1)

corolla carnose, campanulate, 2.8—3.2 mm long, the tube 1.2—1.8 mm long,

glabrous, the lobes ovate, 1.3—1.8 mm miong, 0.8—1 mm wide, erect to spreading,
apically attenuate toa round tip, prominently or conspicously brown punctate,
sparsely lepidote without, densely glandulat-granulose on the upper half
within, the margin entire, glandular-granulose; stamens 2.2—2.6 mm long,
the staminal tube conspicuous, carnose, 1.2—1.6 mm long, lobate, the lobes
to 0.2 mm long, the apically free portions of the filaments 0.4—0.6(—7) mm
long, the anthers triangular, 0.7-0.9 mm long, 0.4—0.5 mm wide, slightly
distally recurved, dehiscent by narrow introrse slits, apically attentuate to
an apiculate tip, basally cordate, the connective inconspicuously brown punctate;
pistillode elongate, conic, 1.2—-1.8 mm long, densely GapsiuceD glandu-
lar-lepidote ay Pistillate inflorescence as in staminate but 1—3 cm long;

floral bracts 1—1.3 mm long, ca. 0.2 mm wide; pedicels 0.6—-0. 9 mm long.
Pistillate flowers as in staminate but calyx ca. 1 mm long, the tube 0.1—0.2
mm long, the lobes 0.8—1 mm long, 0.4—0.6 mm wide; corolla, staminodes
and pistil unknown. Fruit subglobose, 4—6 mm long, 5—7 mm diam., exo-
carp thin, pellucid punctate.

Distribution. —Venezuela, Ecuador, Peru, Amazonian Brazil, and reported
here for the first time from Bolivia, from 70-700 m.

Ecology and conservation status. —Cybianthus gnyanensis subsp. psendoicacoreus
inhabits igap6 forests of South-Central and Western Amazonia. It occurs
in these forests on deep white sands just below the floodline. It is periodi-

cally inundated, but not for long periods. Quantitative fieldwork in Peru
has shown it occurs in populations of 8-10 individuals > 2.5 cm DBH per
hectare.

Etymology.—The subspecific epithet refers to the growth habit of the plant,
somewhat reminiscent of Ardisza (subgenus Icacorea) guyanensis (Aublet) Mez.

R ined. ECUADOR. Napo: Canton peas Reserva Etnica
a aorani, eae Oil Hwy., km 60-61, S of Rfo Tivacuno, 00° 51'S, 76° 26' W, 250 m,

25 Oct 1993 (fr), M. Aulestia & J. Andi 925; Maxus eae pipeline Rd., km 68,
10 a SW . Rfo Tivacuno, 00° 49'S, 76° 26! W, 240 m, 13 Dec 1993 (fr), D. Nez// et al.
10303 (BRIT, MO, QCNE). (BRIT, MO, QCNE); Estacion Experimental INIAP-Payamino,
5 km NE of Coca, 00° 26'S, 77° OL' W, 250 m, 18—26 Feb ae (fr), W. Palacios et al.
1040 (MO, NY, QAME), (stam. fl), W. Palacios et al. 1050 (MO, NY, QAME), 00° 25'S,
77° 00' W, 250 m, 29 Nov 1986 (fr), D. Nei// 7494 (MO, QAM Parque Nacional Yasunf,
Lagunas de Garza Cocha, shore of Rio Garza, 01° 01'S, 75° 47' W, 200 m, 22 Sep 1997
(fr), C. Cerén & N. Gallo 5063 eee QCNE), trail deve the house, 850 m, 27 Apr 1986
ee ee SSOT (QCA). PERU. Loreto: Prov s, Iquitos, G. Bee 3650
(NY), 10 3-11 Aug 1929 is E, Killip G A.C. sn 27 7605 5 (F, NY, US); Mishuyacu,
near oi 100 m, 1930 (stam. fl), G. Klvg 1412 (F), Feb 1932 (fr), o 2565
NY) 1929 (stam. fl), E. Killip & A.C. Smith ae Deto. Alto Nanay, trail ate
N from - a. f Sta. Marfa de Nanay, 5 Mar 1968 (stam. fl.), D. Simpson & J. Schunke 784
(F, US); Deco. Indiana, Explorama Lodge, halfway between Indiana and mouth of Rio Napo,
03° 28'S, 72° 50' W, 130 m, 26 Jun 1983 (fr), A. Gentry et al. 42183 (AMAZ, MO), Far

Prpoty, Cybianthus in Ecuador and Peru pal

end of Bushmaster Trail, 140 m, 5 Jan 1991 (ster.), A. oe et al, 72129 (AMAZ, MO,
US, USM), aa Lodge, near Yanamono, 25 km ae O33 OF S57 22 500M
106 m, 24 Nov 1981 (fr), R. Vdsquer GN. Jaramillo a (AM MO, US, USM), Pe-
rimeter path at ee boundary of reserve, 110 m, 27 Sep 1550 (ster.), J. Pipoly et al.
12492 (AMAZ, MO, US, USM), 03° 28'S, 72° 52' W, 106 m, 15 Apr 1992 (fl bud), R.
Vasquez & N, Jaramillo 18240 (AMAZ, BRIT, MO,USM); Dtto. ee aes yo, ies 6
Experimental del IIAP, 04° 10'S, 73° 30' W, 150-180 m, 3 Nov O (fr), R. Vasquez &
N. Jaramillo 14545 Seta ane MO, USM)8 Nov 1990 (ster.), R. ee N. Jaramillo
15016 (AMAZ, MO, US, M); Puerto Almendras, 2 Jul 1992 (fr), R. Rueda ee Rutz
597 (AMAZ, noes MO); Bee aliaegias Arboretum Colegion Ingeneria Fores Pp,
03° a 73° 25' W, 122 m, 4 Sep 1992 (ster.), C. Grandez et ae ee (AMAZ, ge

O, M); Detto. Las nomen Quebrada cee O32 15! °55' W, 140 m,
Aug i (stam. fl), f ‘tiz et al. 74 (AMA RIT, MO, cae Dtto. Sta. Maria _
Nanay, wpiees 03° ie 73° 35' W, 90 . = 1990 (ster.), J. Pipoly et al. 12706
(AMAZ, MO, USM). Madre de Dios: i ee: Cerro de Pantiacolla, Rio Palotoa, 10—
15 km NNW of eee 12° 35'S,71° 18' W, 650-700 m, 13 Dec 1985 (ster.), R. ee
et al. 10993 (FE, NY, USM); Prov. a Santuario Nacional Pampas del Heath, Rio

Heath, 12° 39! 23"S, 68° 44' 13" W, 210 m, 5 Jun 1996 (stam. fl), M. Aguilar & D. Castro
cee (BRIT, MO, USM). BOLIVIA. Santa Cruz: Dtto. Velasco, Parque Nacional Noel
Kempff Mercado, Campamento La Torre, 13° 39' 20" S, 60° 49! 08" W, 200 m, 24 Nov
1993 (fr), L. Arroyo et al. 510 (BRIT, MO, USZ).

Cybianthus guyanensis subsp. pseadoicacoreus is one of three subspecies. The
species is defined by the autapomorphic contorted anthers of the stamens
and staminodes (Pipoly 1992a). The three subspecies may be separated in
the following key.

1. Leaf blades symmetric; calyx subcupuliform; corolla chartaceous or carnose,
the lobes erect to spreading; apically free portions of the filaments shorter

than the anthers; anthers narrowly triangular or ovate-triangular, apically
apiculate, dehiscent by narrow, introrse slits
etioles canaliculate and winged, (1—)1.5—1.9(—2.3) cm long; staminate
peduncle 0.2—0.5 cm long; fae bracts shorter than the pedicels, 0.7—
0.8 mm long; pedicels 0.5—1 mm long; corolla chartaceous, 2.3—2.6 mm
long, the lobes elliptic; ane eae 0.9- 1.1 mm long; apically free portions

of the filaments 0.2—0.4 mm oe subsp. guyanensis
2. Petioles canaliculate, 0. ne 1.5) cm long; staminate peduncle 0.1—0.2

cm ee Wo ibiaer 1onees than the pedicels, 1.3—1.5 mm long; pedicels

1—1.2 mm long; corolla carnose, 2.8—3.2 mm long, the lobes ovate; staminal

tube 1.2-1.6 mm long; apically fee portions of the filaments 0.4—0.
(7) mm long subs] pee i ueiecleus
the

—

. Leaf blades asymmetric; calyx cotyliform; corolla oe ae
lobes reflexed-recurved; apically free portions of the filaments longer than
the anthers; anthers ovate, apically acute, dehiscent by wide, sublatrorse slits.
subsp. multipunctatus

Cybianthus subspecies multipunctatus (A. DC.) Pipoly is distributed in eastern
Amazonia and the Guianas in premontane forests on lateritic and white sands
of the Roraima Superimposed Sediments, while subsp. gwyanenszs is located
principally in central Amazonia in igapé forests (Pipoly 1992a). Subspe-

52 Sipa 18(1)

Steps f? Ries sil

Fic. 13. Cybianthus timanae Pipoly. A. Habit, showing paniculate inflorescences. B. Branchlet
apex. C. Adaxial leaf surface, showing midrib slightly raised but canaliculate. D. Abaxial
leaf surface, showing lepidote scales and prominently raised midrib. E. Section of raceme
showing coriaceous, deltate floral bracts, cotylitorm calices, obcordate anthers with dis-

tally recurved apiculae. F. Pistillate flower bud. G. Pistillace corolla with one lobe removed,
showing en pistil and subsessile antherodes. A-E, drawn from holotype. F—G, drawn

from Timand 1047. Figure drawn by Linda Ellis.

Prpoty, Cybianthus in Ecuador and Peru a3

cies pseudoicacoreus is distributed south-central and western Amazonia, along
banks of smaller streams on deep white sands. Subspecies psewdoicacoreus and
guyanensis are sympatric only in central Brazilian Amazonia, and differences
in their ecology are not known in sufficient detail. In Ecuador and Peru,
subspecies psewdoicacoreus may be most easily confused with C. comperuvianus
Pipoly, but may be recognized by the thicker branchlets, longer floral bracts,
and carnose perianth.

10. ip anaeaes timanae Pipoly, _ nov. (Fig. 13). Type: PERU. JuNiN: Prov. Satipo,
onal, Mapari, ca. 12 km SW of Chequitavo, 10° 45'S, 74° 23' W, 1,300 m,
7 Apr oe (stam. fl), D. Smith . (HOLOTYPE: MO; tsoryPes: BRIT, US, USM).

no ram es graciles ae laminas sence a Opiate ad apices acuminatas
- bi

minaret inconspic uo, C. occigranatensi arcte affinis, sed ab ea nerviis secundariis
16—40 (non 8-12)-} jugis, perianthiis chartaceis (non coriaceis), corolla coo oun cca

eae i lobi llinis laevibus (non verrucosis)

conspicue (nec incounspicue) atro-punctatis, pistilo laseniform (nec obnapiformi) perfacile
separabilis.

Subshrub to 1 m tall. Branchlets angulate, 2—2.5 mm diam., densely lepi-
dote. Leaves alternate; blades chartaceous, elliptic to oblanceolate, (8—)
9.5—15(—19) cm long, (2.7-)3—-5.5(-6.5) cm wide, apically abruptly acuminate,
caudate, the acumen 1—2.2 cm long, basally acute, decurrent on the peti-
ole, smooth and nitid above, pallid and moderately lepidote below, midrib
slightly raised and canaliculate above, not decurrent on the petiole, promi-
nently raised below, secondary veins 16-40, brochidodromous, planar to
somewhat impressed above, not bullate, the margin flat, entire; petioles
canaliculate, (1.3—)1.5—2(—2.2) cm long, glabrous above, densely lepidote
below. Staminate inflorescence: a panicle of 2-8 racemes branched from base,
(3—)4—6(—7) cm long; peduncle 3—7 mm long; rachis densely lepidote; floral
bracts coriaceous, deltate, 0.6—0.8 mm long and wide, apically acute, somewhat
cucullate, densely lepidote above and below, the margin entire; pedicels
cylindrical, 1-2.5 mm long, densely lepidote. Staminate flowers 4-merous,
cream; calyx chartaceous, cotyliform, 0.7—1 mm long, the tube ca. 0.2—0.3
mm long, the lobes ovate-triangular, 0.5—0.7 mm long, 0.3—0.4 mm wide,
apically acute, moderately lepidote, the margin entire, somewhat involute;
corolla chartaceous, cotyliform, 2—2.4 mm long, the tube ca. 0.2 mm long,
the lobes oblanceolate, 1.8—2.2 mm long, 0.8—1 mm wide, apically subacuminate,
sparsely lepidote apically near margin, prominently black punctate and punctate-
lineate without, densely glandular-granulose throughout within, the mar-
gin glabrous, entire; stamens 1.4—1.6 mm long, the tube conspicuous,
chartaceous, 0.1—0.2 mm long, elobate, the apically free portions of the filaments
terete, 0.4—0.6 mm long, the anthers obcordate, 0.6—0.7 mm long, 0.3—
0.4 mm wide, apically apiculate, anther and apiculum distally recurved,

54 Sipa 181)

the connective prominently black punctate dorsally; pistillode conic, 1.2—
1.4 mm long, 0.3—0.4 mm wide, hollow, densely translucent glandular-
lepidote. Pistillate inflorescence as in staminate, but (1—)1.5—3 cm long; pe-
duncle 2-4 mm long; floral bracts 0.4-0.6 mm long and wide; pedicels
obconic, 0.6-0.9 mm long, sparsely lepidote, conspicuously black punc-
tate. Pisti/late flowers as in staminate but translucent green; calyx 0.7—0.9
mm long, the tube ca. 0.2 mm long, the lobes 0.5—0.7 mm long, 0.3—0.4
mm wide, sparsely lepidote, corolla 1.2—1.5 mm long, the tube ca. 0.2 mm
long, the lobes ob! long to oblanceolate, 0.9-1.3 mm long, 0.4—0.6 mm wide,
apically obtuse to subacuminate; staminodes 0.6—0.8 mm long, the tube
conspicuous, chartaceous, ca. 0.1 mm long, the antherodes subsessile, 0.6—
0.7 mm long, 0.3—0.4 mm wide; pistil lageniform, 1.2—1.4 mm long, 0.3—
0. . mm wide, hollow, densely translucent glandular-lepidote, the ovules
2—3, partially immersed on the placenta. Frvit slightly depressed-globose,
4, it 5.5 mm long, 5.5—7.5 mm diam., the exocarp thin, black at maturity.

Dish but on Sontheascetn Ecuador to Cusco Peru, at 720—1,300 m
elevation.

Ecology and conservation status. —Cybianthus timanae usually occurs in wet
premontane forest on sandstone soils. The restricted distribution of Cybianthus
timanae indicates it should be considered a threatened species.

Etymology.—The species is named for Martin Timana de la Flor, former
Peruvian Field Associate of the Missouri Botanical Garden, and currently a
graduate student at the University of Texas at Austin. Martin is specializ-
ing in the systematics of high altitude Caryophyllaceae.

PARATYPES, ECUADOR. Zamora-Chinchipe: Nangaritza poe Owe hee of
Cordillera del Céndor, above Pachicutza, Rio Nangaritza Valley, 04
1,000-1,200 m, 6 Dec 1990 (fr), D. Neill & W. Palacios 9556 pee MO, Qc NF Ri

pete Shaime, confluence of Rios Nangaritza and Numpatakaime, 04° 20! S,
'W, 1,000 m, 7 Dec 1990 Why . tee au (BRIT, MO, . oie a. ie to
i. aie del Condor, 04 , 78° 37! W, 1,000-1,10 19 Oct 1991 (piste. fl

bud), W. Palacios et al. 8: ae a Nie OCND), 20 Oct 1 a coon fl), W. Palacios
et al. 8407 (BRIT, COL, MO, PORT, QCNE, USM); Say ee NE of military
camp, 900 m, 6 Dec 1990 (fr), J. Jaramillo & E. Grijalva 13419 (COL, QC pene
oe narca: Cutervo National Park, 12 km NE of San Andrés de Cutervo, Transect 3, 06°
10'S, 78° 40! a 2,230 m, 10 Sep 1991 (ster.), A. Gentry et al. 74630 (BRIT, ve cour
Cusco: Prov. Quispicanchi, Camanti, Maniri, along trail parallel to Rfo Maniri to Quebrada
Garrote, 13°17’S, 70°48’ W, 720 m, 17 Oct 1990 (pist. fl), M. Timand 1047 (BRIT, CUZ
MO, US, USM)

Cybianthus timanae is most closely related to C. occigranatensis (Cuatrec.)
G. Agostini, but may be easily separated by its more numerous secondary
veins, chartaceous perianth parts, cotyliform corolla with smooth, conspicuously
black punctate, acuminate lobes, and lageniform pistil. The involute calyx
lobes and the obconic pedicels of the pistillate flowers are also extremely
rare within the subgenus.

Prpoty, Cybianthus in Ecuador and Peru 55

1 - s ]

Fic. 14. ee Pipoly. A. Habit shaped, basally branched
panicles. B. Staminate flower, habit, showing cotyliform ealiese infundibuliform corolla.

Opened staminate flower, on owing inconspicuous staminal tube and lageniform aecltcees
D. Opened pistillate ene showing obnapiform pistil and subsessile antherodes. A—C,
drawn from holotype. D, drawn from A. Gentry et al, 53960. Figure drawn by Juan Pinzén.

(le gone Sorina Pipoly (Fig. 14), Caldasia 18(3):285. 1996.

Huita: Municipio La Plata, Vereda Agua Bonita, Finca Merenberg,

cerca r eens Gardin Bocinice! 22 Abr 1982 (stam. fl) J. H. Torres 1054
(HOLOTYPE: COL)

Dioecious sree to 6 m tall. Branchlets narrowly angulate, 3-4 mm diam.,

56 Sipa 18(1)

densely appressed-lepidote, the scale margins frequently overlapping. Leaves
alternate; blades chartaceous, elliptic or oblanceolate, 8-20 cm long, 3.5—
6.5(—6.9) cm wide, apically acuminate, the acumen (0.5—)1.0—1.5(—2.0) cm
long, basally acute, slightly decurrent on the petiole, midrib impressed above,
prominently raised below, the secondary veins 26—34 pairs, inconspicuous
above, prominently raised below, sparsely lepidote above at first, then pus-
tulate and glabrescent, densely appressed-lepidote below with margins fimbriate
and overlapping, the blade margin flat, entire; petioles canaliculate, (1.0)1.3—
1.7(—2.0) cm long, densely appressed-lepidote. Staminate inflorescence: an irregular
panicle with 2—4 basal branches, 4-8 cm long; peduncle, rachis and pedicels
densely lepidote; peduncle 0.1—0.4 cm long; floral bracts chartaceous, tri-
angular, subequalling the pedicels, 1.2-1.5 mm long, ca. 0.6 mm wide,
apically attentuate, densely lepidote below, the margin entire, glabrous; pedicels
cylindric, 1.0-1.5 mm long. Staminate flowers 4-merous; calyx carnose, cotyliform,
1.3-1.4 mm long, the tube 0.2—0.4 mm long, the lobes triangular or ovate-
triangular, 1.0—1.2 mm long, 0.6—0.9 mm wide, apically attenuate, con-

spicuously brown punctate, the margin entire, regular, with few scales; corolla
chartaceous, infundibuliform, 3.0—3.5 mm long, the tube 1.2—1.3 mm long
the lobes ovate, 2.0—2.2 mm long, 1.0-1.2 mm wide, apically rounded,
carinate medially, densely lepidote without and along the margins, glan-
dular-granulose toward the apex and along the margins within, inconspicuously
brown punctate, the margins regular, entire; stamens 2.5—3 mm long, the
staminal tube membranaceous, inconspicuous, 1.2—1.3 mm long, hyaline,
elobate, glabrous, the apically distinct filaments adnate to the corolla lobe
above the staminal tube 0.3—0.5 mm and apically free 0.1—0.3 mm, gla-
brous, the anthers ovate, 0.8—1.0 mm long, 0.5—0.6 mm wide, distally reflexed,
apically apiculate, the ae clans slightly inflexed, basally cordate, the con-
nective dark, eglandular; pistillode lageniform, 1.6—-1.8 mm long, densely
glandular- nepicote basally. Pisti//ate inflorescence: as in the staminate except
a simple raceme, 4.4-10 cm long; ponace 0.2—0.5 cm long; floral bracts
lanceolate, 1.2—1. ce long, 0.3-0.4 mm wide; pedicels 0.7—-1.3 mm long,
incrassate in fruit. Prstz//ate flowers similar to the staminate except calyx 1.4—
1.7 mm long, the tube 0.2—0.4 mm long, the lobes ovate, 1.3—1.5 mm long,
0.6-1.2 mm wide; corolla 3.5—4.0 mm long, the tube 0.9-1.1 mm long,
the lobes elliptic, 2.7—3.1 mm long, 0.9—-1.3 mm wide, apically acute; staminodes
similar to stamens but 3.0—3.2 mm long, the staminodial tube 1.0 mm
long, the apically free portion of the filaments adnate to corolla lobes 1.3—
1.5 mm and apically free ca. 0.1 mm, the antherodes subdeltate, 0.8—1.1
mm long and wide, (always longer than wide), apically acute, basally sag-
ittate; pistil obnapiform, 1.8—2.0 mm long, the ovary 0.9-1.0 mm long,
1.2—1.5 mm diam., densely translucent glandular-lepidote, the style 0.9—
ong, the stigma bilobate, not capitate, the placenta deeply cupuliform,

_—

1.1 mm

Pipoty, Cybianthus in Ecuador and Peru 57

the ovules 2, deeply embedded in the placenta below apical pores. Fruit
drupaceous, depressed-globose, 4-6 mm long, 6-8 mm diam., the endocarp
smooth, the embryo erect, 3 mm long.

Distribution. —C ybianthus cuatrecasasii occurs in the Western Cordillera
of Colombia in the Department of Antioquia, in the Central Cordillera near
the Macizo Colombiano, in the Departments of Huila and Cauca, and in
the Western Cordillera of adjacent Ecuador, with disjunct populations in
the Province of Santiago-Zamora, and Zamora-Chinchipe, Ecuador, from
(1,200—)1,960—2,850 m elevation.

Ecology and conservation status. —Cybianthus cuatrecasasii occurs along the
margins of primary cloud forests and along small watercourses. The popu-
lation from Antioquia, Colombia is rare because it is from premontane plu-
vial forest, and has leaves much shorter than normal. Because it is known
only from primary forests, it should be considered threatened.

Etymology.—This species is named to honor the late José Cuatrecasas, prodigious
field botanist, monographer of many plant families, and ardent student of
the Colombian flora. Don José freely offered his advice and assistance to all
who asked, despite his limited time and numerous projects.

Specimens examined. COLOMBIA. Antioquia: Mpio. San Luis, Piedra de Cascrill6n,
3—4 hours by foot S of 06° O1'N, 75° OL! W, 1,500-1,700 m, 8 May 1989 (fl bud),
D. Daly et al. 5926 CHUA, MO, NY, US). Huila: Cordillera Central, E slope, Finca Merenberg,
km 101 of La Plata-Popayén Rd., 13 km E of Sta. Leticia, 02° 15'N, 76° 12' W, 2,300 m,
24 Mar 1986 (fl bud), B. Stein 3721 (BRIT, MO); Cordillera entre cuencas de los rfos Guarapas
y Guachicas, arriba de Palestina, al SW de Pitalito, 2,000—2,300 m, 6 Feb 1943 (stam. fl),
ER. Fosberg oe (NY, US); Finca Merenberg, E de Volcan Puracé, cerca de la zona limitrofe
con Cauca, 26' N, 76° 12' W, 2,300 m, 1 Apr 1986 (bud), A. Gentry et al. 55881
(COL, MO, oS 3 Apr 1986 (pist. fl, fr), A. Gentry et al. 53960 (COL, MO, US); 15 km
NE de Algeciras, cerca del Campamento La Gironda, 2,400-2,850 m, 26 Mar 1944 (stam.
fl) E. Little 7481 (COL, NY, US); Mpio. San José de Isnos, Vereda El Hornito, 1,960—2,000
m, 23 Jul 1980 (stam. fl), G. Lozano 3366 (COL); Mpio. La Argentina, arriba de Finca
Palmira, 2,100 m, 26 Sep 1984 (fl bud), G. Lozano et al. 4133 (COL). Cauca: Moscopan,
margen del Réo San José, La Chorrera de Candelaria, 2,100—2,350 m, 1 Feb 1947 (fr), J.
Cuatrecasas 23583 (COL, F, NY, US); Cuenca del R fo La Plata, 43 km SE de Popayan, 2,160
m, 26 Nov 1944 (stam. fl), ER. Fosberg 22376 (NY, US). ECUADOR. Santiago-Zamora:
along Quebrada Honda, vicinity of Rancho Achupallas, 2,500—2,700 m, 10 Oct 1943 (stam.
fl), J. Steyermark 54552 (NY). Zamora-Chinchipe: Hill ca. 2 km downstream from Campamento
Shaime along Rio Nangaritza, 900 m, 15 Feb 1994 (fr), H. van der Werff 13074 (BRIT,
MO, QCNE)

Cybianthus cuatrescasasii is unique within subgenus Conomorpha because
of its infundibuliform corolla and subapical portions of the filaments ad-
nate to the corolla lobe. The overlapping covering scales of the abaxial leaf
surface, branchlets and inflorescence rachis is found otherwise only in Cybianthus
crotonoides (Mez) G. Agostini, a species endemic to the Guayana Highland.
This species has been confused with Cybianthus peruvianus (Mez) G. Agostini,

58 SIDA L8(1)

but is easily separated from it by the infundibuliform corolla, long pedicels
and eglandular connectives. Cybianthus cnatrecasasil 1s Closely related to C.
laetus (Mez) G. Agostini and C. oceigranatensis (Cuatrec.) G. Agostini, from
which it is easily distinguished by its infundibuliform corolla, apical por-
tions of the filaments adnate to the corolla and lack of lobes on the staminal
and staminodial tubes.

Hil. Pe subgenus Laxiflorus G. Agostini, Acta Biol. Venez. 10:144.
1980; Pipoly, Bricttonia 35:61—80. 1983. Conomorpha section Conomorphida Migq.,
Stirp. Surinam Select. 111. 1851, ye ee Cipnanipia St subgenus Ewe deca ees M
in Engl., Pflanzenr. Iv. 236(Heft 9):254. 1902. j TYPE SPEC ee
laxiflora (Mart.) A. DC. = C eed os (H. B. K.) G. Agostini. ce TOTYPE: by
Agostini, Acta Biol. Venez. 10:144. 1980)

Terrestrial dioecious shribs or small trees. Roots positively geotropic. Trunk
distinguishable, leptocaulous, the growth dynamics following Rauh’s Ar-
chitectural Model (Hallé et al. 1978). Branchlets tomentose, with dendroid
or stellate ferrugineous trichomes or both. Cataphy//s and pseudocataphylls
absent. Leaves alternate, petiolate, tomentose, glabrescent. lnflorescence a simple
raceme or rarely a spike, the peduncle 0.1—0.5 cm long, the staminate ra-
chis tortuous or lax, the pistillate rigid and erect or rarely lax. F/owers 4- or

5-merous; calyx cotyliform, valvate, epunctate, inconspicuously pellucid or

conspicuously brown or Orange punctate, glabrous, tomentose, or g landu-
lar-granulose, rarely bearing translucent glandular scales, the margins glandular-
ciliate; corolla campanulate, the lobes valvate, glandular-granulose only along
margins without, but over the entire surface within, epunctate or incon-
spicuously pellucid or conspicuously brown or orange punctate, and punc-
tate-lineate, medially and basally glabrous, glandular-granulose or with a
few stellate trichomes; stamens and staminodes with filaments basally con-
nate into a conspicuous or inconspicuous tube, the tube elobate or bearing
small lobes alternate with the apically free filaments, the anthers elongate,
triangular, distally curved, rarely erect, dorsifixed 1/4—1/2 from base, apically
acute, basally cordate, dehiscent by wide longitudinal slits; staminodes re-
sembling stamens but reduced in size, the sterile anthers without pollen or
at times producing abortive pollen; pistil obnapiform, the ovary translu-
cent glandular-lepidote, the style short, truncate with a punctiform stigma;
pistillode lageniform, the ovary translucent glandular-lepidote, hollow the
style elongate and curved apically, the nonfunctional stigma capitate. Frit
drupaceous, one-seeded, the exocarp thin.
Cybianthus subgenus Laxiflorus contains 6 species, 3 of which have been
recorded from Peru. To date, none are known from Ecuador, but we may
expect Cybianthus spicatus ISunth) G. Agostini in sandstone areas in the Cordillera

del Céndor area.

Pipoty, Cybianthus in Ecuador and Peru 59
KEY TO SPECIES OF CYBIANTHUS SUBGENUS LAXIFLORUS

1. Inflorescence rachis black PUBCArS, calyx lobes coriaceous, deltate, medi-

“4

A

1)

a
5

g

S

jon
iw]

4

ct
a
=

1)

a
°
5
Et io)

i)
a
ite)
oO

ally carinate, basally rugose globose,
trees to 18 m tall, serra ne Pilla seco” [dry, on white sand} forests.
12. C. nestorii

1. Inflorescence rachis epunctate; calyx lobes pale ce widely to narrowly
ovate-triangular, medially thicked but flat; basally smooth; fruit globose, as
broad as long; shrubs or small trees to LO(—12) m a riparian gallery for-
ests, either eerie flooded (varzea or igap6) or “varillal hamedo” [wet,
white or brown sand ests
2. Leaf blades chartaceous, the tertiary veins prominently raised above; ca-

lyx lobes longer than broad; staminal and staminodial tubes inconspicu-
ous; receptacle or pedicel or both enlgarged in fruit; seasonally inundated
13. C. spicatus

forests (varzea or igap6)
2. Leaf blades coriaceous, the tertiary veins not visible from above; calyx
lobes broader than long; sages and staminodial tubes conspicuous;
ither receptacle nor pedicel enlarged in fruit; forests with
poor ine on sand, but ee nee Cwatillal bumedo )ice: 5.
14. C. fulvopulverulentus subsp. magnoliifolius

12. Cybianthus nestorii Pipoly, sp. nov. (Fig. 15). Type: PERU. Lorero: Prov.
Maynas, Mishana, on Rio Nee 03° 53'S, 73> 32 WW, 15 Orm:, 20: Jam 10a 5;
R. Vasquez & N. “Tavamillo 6122 (HoLoryPE: MO; isotypes: AMAZ, F, NY, US, USM).

e delrar.

Ob folia coriacea, rhachides inflorescentiares atro-punctato-lineatos,
inter se aequilatos coriaceosque, C. de/tato valde arcte affinis sed ab ea Fane arboreo (non
fruticoso) usque ad 18 (non 3) m, laminis anguste ellipticis vel anguste oblanceolatis vel
anguste oblongis (non habia 9.5-18(—28), (nec 3.2—4.4) cm longis, 3.2-4.6 (nec 1.7—
2.8) cm latis, inflorescentiis 5—7 (non 1—4.2) cm longis, lobis calycinis 1.8—2 (non 1.3—
1.7) mm longis, eet (nec planisque) denique secus marginem minute erosis (nec
undulatis), statim separabilis.

Tree to 18 m tall. Branchlets terete, S—7(-15) mm diam., appressed fer-
rugineous dendroid tomentose. Leaves alternate, blades thinly coriaceous,
narrowly elliptic to narrowly oblanceolate, rarely narrowly oblong, (9.5—)
11.5—14(-18) cm long, (3.2—)4—6 cm wide, apically obtuse, emarginate,
rounded or acute, basally acute to attenuate, decurrent on the petiole, gla-
brous and nitid above, glabrous and nitid but pallid below, midrib slightly
raised above, prominently raised below, decurrent through length of peti-
ole above and below, the secondary veins (14—)20—47 pairs, prominulous
(slightly raised) above and below, inconspicuously pellucid punctate and
punctate lineate below, the margin entire, glabrous, revolute; petioles mar-
ginate, (2.2—)2.5—3.2(-3.7) cm long, glabrous. Staminate inflorescence: un-
known. Pistillate inflorescence: an erect raceme, S—7 cm long, the rachis black
punctate-lineate, ferrugineous glandular-granulose, glabrescent; floral bracts
coriaceous, linear, minute, 1—1.1 mm long, 0.2—0.3 mm wide, apically acute,
densely glandular-granulose below, glabrescent, the margin glandular-cili-

60 Sipa 18(1)

Fic. 15. Cybianthus nestorit Pipoly bose fruits and leaf blades
with prominulous secondary veins and revolute margins. B. Portion of infructescence, showing
depressed-globose fruit and carinate, rugose calyx lobes with erose, sparsely glandular-ciliolate
margins. C. Abaxial leaf surface, showing prominently raised midrib and prominulous secondary
veins. D. Branchlet apex, showing appressed dendroid tomentum. A—D, drawn from holo-

type, by Linda Ellis.

_ A. Habit, showing depressed-globose fr

Pipoty, Cybianthus in Ecuador and Peru 61

ate; fruiting pedicels cylindrical, 24 mm long, densely glandular-granulose,
minutely black punctate apically. Pést///ate flowers unknown; fruiting calyx
coriaceous, 1.8—2 mm long, the tube 0.3—0.5 mm long, the lobes deltate,

1.3-1.6 mm long, 1.4-1.7 mm wide, apically acute, medially carintae, rugose
basally, the margin minutely erose, sparsely glandular-ciliolate; corolla, staminodes
and pistil unknown. Fruit depressed-globose, 4—6 mm long, 6-8 mm diam.,
black at maturity, inconspicuously pellucid punctate.

Distribution.—Known only from the Rio Nanay Drainage Basin, Maynas
Province, Department of Loreto, Peru, at 150 m elevation.

Ecology and conservation status. —Cybianthus nestorii occurs in “varillal seco”
habitats, consisting of tall terra firme (non-inudating), relatively dry forest
on deep white sands. It is a rare species and thus, should be considered threatened.

Etymology.—It is a pleasure to dedicate this species to Nestor Jaramillo,
of Iquitos, Peru, prodigious collector, and authority on plant collecting in
tropical forests. Nestor, with his supervisor, Rodolfo Vasquez, form one of
the most formidable botanical exploration teams in South American botany.

Paratypes: PERU. Loreto: Prov. Maynas, Dtto. Iquitos, Puerto Almendras, UNAP, Tree
No. 324, Tree inventory, 03° 48'S, 73° 25' W, 122 m, 17 Jan 1993 (ster.), C. Grandez, N
Jaramillo et al. 5321 (BRIT, MO, UNAP), Tree No. 373 (ster.), C. Grdndez, N. Jaramillo et
al, 5370 (BRIT, MO, UNAP, USM); Tree No. 651 (ster.), C. Grdandez, N. Jaramillo et al.
5642 (BRIT, MO, UNAP)

Cybianthus nestorii is unique within the subgenus because of its autapomorphic
depressed-globose fruits and carinate calyx lobes. Other characters which
readily seaparate it from its closest relative, Cybzanthus deltatus Pipoly, of
the Rio Guainia drainage basin of Venezuela, include its much larger ar-
borescent habit, larger elliptic to narrowly oblanceolate leaves, longer
inflorescences, and longer, minutely erose calyx lobes.

13. Cybianthus spicatus (Kunth) G. eee (Fig. 3B), Acta Biol. Venez.
10:146. 1980. Myrsine spicata Kunth in H.B.K., Nov. Gen. Sp. 3:250. 1818. C —
Nes (Kunth) Mez in Engl., Pflanzenr. = 236(Hef 9):259. 1902. Typ

“UELA. Terrirorio FEDERAL AMAZONAS: In inundated area along the Rio ae oe
nee date ae fl), A. von Humboldt & A, ae: 1096 (HOLOTYPE: P-B
ISOTYPE: B-d

sta meen Mart., Nov. Gen. Sp. Pl. 3:89. 1829. ue laxiflora (Mart.) A.
Soc: London, Bot. 17: 102. 1834. T . AMAZONAS: Prov.
a Nestor Th pee eae Jan 1826 on ae a ee 5.2. (LECTOTYPE

y Pipoly 1983b: M; IsoLecroryPe: E).

Common laxiflora vat. ln Miq. in Mart., Fl. Bras. 10:302. 1856. Type: BRA-
ZIL. AMAzonas: Prov. Rio Negro, vicinity Bier eee Dec—Mar 1850-51 (stam.
fl), R. ae 1040 ee roryee, here designated: M; isoLecroryprs, CGE, GH, GOET,

LY OXF,

om

[soy
Conomorpha laxiflora var. latifolia Mig. in Mart., Fl. Bras. 10:303. 1856. C si ie sal
(Mig. in Mart.) Mez in Engl., Pflanzenr. IV. 236(Heft 9):255. 1902. Typr: VEN

62 Sipa L8(1)

[GUYANA]. 1839 (stam. fl), R. Schomburgk 1002 (LeCroryre by Pipoly 1983b: G;
ISOLECTOTYPE: CGE
Conomorpha a ee Mez in Engl., Pllanzenr. IV. 236(Heft 9): 256. 1902. Type: VEN-
EZUELA. [GUYANA]. 1840 (stam. fl), R. Schomburgk 885 (HOLOTYPE: B-destroyed;
ragment at F; tecroryps, here designated: G; isotecrorypes: CGE, GH, K-2 sheets,

=
~

US, W).
eae grandiflora Mez in Engl., Pflanzenr. 236(Hett 9):258. 1902. Type:
BRAZIL.AMAzoNas: Rio Negro, Gece Moureira, a 1851 (stam. fl.), R. Spruce 1946

(HOLOTYPE: B- serve LECTOTYPE, here designated: K
G ae i bees a, 8 1

C ees Engl., ee - _ 9): 260. 1902. Type: BRAZIL.
oe AS! Rio Negro, about 00° 30'S, 64° 00! W, 24 Jun 1874 (stam. fl), J. Trail
SOS (HOLOTYPE:

Conomorpha madeirensis AC . Smith, J. Arnold Arbor. 20:300. 1931. Type: BRAZIL. AMAZONas:
I moe of Humayeé, between Monte Cristo and Sta. Victoria on Rio ae
15-17 934 (stam. fl), B. Krakoff i (HOLOTYPE: NY; isotypes: A, FM, MAD-

’, MICH “MO, S, US).

G sie ti wal A.C. Smith, Bull. Torrey Bot. Club 67:295. 1940. Typr: GUYANA.
ubaru River, Kako Territory, Upper Mazaruni drainage, 600 m, 2 Feb 1939 (stam.

), A. Pinkus 181 (HOLOTYPE: NY; isotypes: BR, F, GH, M, MO, NY, S, US).

» ISOLECTOTYPES: CGE, GI

Shrub or small tree to 12 m tall. Branchlets thin to moderately thick, (3.5—)
—8(—20) mm diam., appressed to foccose-dendroid and stellate ferrugin-
eous tomentose, the stellate hairs often appearing furfuraceous, glabrescent
Leaves alternate; blades thinly coriaceous to char Caceous, to membranaceous,
ovate, elliptic, obovate or rarely oblanceolate, (4.5—)6—-17.5(—21) cm long,
2.9—8(—11.5) cm wide, apically acute, rounded, obtuse or rarely emargin-
ate, basally acute to cuneate, glabrous, pustulate and often nitid above, sparsely
ferrugineous puberulent and minutely glandular-lepidote below, the mid-
rib, secondary and tertiary veins prominently raised above and below, the
midrib decurrent to the base of the petiole, pellucid or black punctations
obscure below, the margin slightly revolute, entire; petioles distinctly marginate,
(1—)1.5—3.2(-4) cm long, stellate and dendroid ferrugineous tomentose, early
glabrescent. Staminate inflorescence: a simple raceme, lax, (3—)6—9(—12.5) cm
long, the rachis epunctate, glandular-granulose with scattered stellate and
dendroid ferrugineous trichomes, glabrate; floral bracts chartaceous, linear-
lanceolate, 0.7—1.7(—2.2) mm long, densely ferrugineous tomentose, early
caducous; pedicels cylindrical, 0.9—2.1(—3) mm long, glabrous to densely
glandular-granulose. Staminate flowers (4A—)S-merous, chartaceous, cream to
yellow, with a sweet odor; calyx shallowly cotyliform, 1.5—2.5 mm long,
the cube 0.3—0.6 mm long, glandular-granulose at first, glabrescent, the
lobes ovate or ovate-triangular, (1.2—)1.4—2 mm long, 0.9-1.3 mm wide,
acute to abruptly acuminate apically, somewhat cordate basally, prominently
thickened medially, conspicuously punctate, the margins mostly entire, a
occasionally erose apically, glandular-ciliate; corolla campanulate, 3.6—4
mm long, the tube 1—1.2 mm long, the lobes ovate, 2.1—3.4 mm ioe -

Pipoty, Cybianthus in Ecuador and Peru 63

2.6 mm wide, See reflexed at anthesis, obtuse to acute apically, epunctate
or conspicuously brown punctate, stamens 1.6-2.2 mm long, the tube

membranaceous, inconspicuous, |—1.5 mm long, the apically free portions
of the filaments 0.3—-0.55 mm long, the anthers elongate-triangular, 0.7—
1.2 mm long, 0.2—0.3 mm wide, dorsifixed ca. 1/3 from base, apically acute,
basally cordate, distally recurved, the connective inconspicuously brown punctate;
pistillode lageniform, 1.3—1.8 mm long, the ovary 0.6—0.9 mm long, 0.5—
0.75 mm diam., densely translucent glandular-lepidote, the style elongate,
0.8—1.3 mm long, slightly curved apically, the stigma capitate, 0.1—0.2 mm
long. Pistillate inflorescence as in staminate, erect, (3.3—)4.5—13(-17.5) cm
long; floral bracts 0.5—2.4 mm long; pedicels 1.3—1.8 mm long, accrescent
in fruit to 3.1 mm long, the receptacle or pedicel or both incrassate in fruit.
Pistillate flowers as in staminate but chartaceous, yellow; calyx cotyliform,
1.3-1.9 mm long, the tube 0.6—0.9 mm long, the lobes widely to narrowly
ovate, 1.1—1.7 mm long, 0.9-1.5 mm wide, erect, sparsely brown punc-
tate, prominently thickened below, the margins entire or occasionally erose
apically; corolla 2.7—-3.5 mm long, the tube 0.7—0.9 mm long, the lobes
1.6—2.7 mm long; staminodes as in stamens but 1.6—2.1 mm long, the tube
chartaceous, conspicuous, 0.7—1.1 mm long, the apically free portions of
the filaments 0.3—0.5 mm long, the antherodes 0.6—0.9 mm long, 0.2—0.3
mm wide, at times producing abortive pollen; pistil obnapiform, 1.6—2.2
mm long, the ovary 0.8-1.2 mm long, 0.9-1.2 mm diam., densely trans-
lucent glandular-lepidote, the placenta cupuliform, ovules 4, partially im-
bedded, the stigma capitate, 0.1—-0.2 mm long. Fruit globose, 2.5—6 mm
long and in diam., prominently black punctate-lineate, with a few persis-
tent lepidote glandular scales below the usually persistent style base, green
then purple, then blac

Distribution.—Venezuela, Colombia, Peru, Brazil and Guyana, in inun-
dated forests ere and igap6), from 100-400 m.

Ecology and conservation status. —C ybianthus spicatus is restricted to primary
riparian habitats on white sandy soils. It is common in igap6 and varzea,
but the latter only when sufficient quantities of sand exist, a mixed varzea
type. While it is a widespread species, it is sensitive to soil compaction,

and should be considered threatened.
Etymology.—The epithet refers to the inflorescence shape, which is a raceme
bearing flowers on short pedicels, thus appearing spicate.

Representative ana examined. PERU. Hudnuco: Cerros del Sira, SW oes of
the Rfo Lulla Pichis, 1190 m, 12 Jul. 1969 (fr), J. Wolffe 12295 (F, NA); 100 m, 22 Jul
1969 (fr), 7; Dudley 131 _ (F, NA); Prov. Pachitea, region of Pucallpa, W part of Sira Mountains
and adjacent lowland, ca 24 km SE to 26 km ESE of Puerto Inca, next to Campament«
Pato Rojo, 09° 27'S, 74° 46' W, 1,000 m, oo 988 (pist. fl), We Morawetz & B. Wallnifer
12-27188 (BRIT, MO, W, WU), 1,320 m, 1 May 1988 (fr), B. Wallnifer 111-1588 (BRIT,
W, WU), 1,230 m, 13 Jun 1988 (fr), B. Wal/ndfer 1112-13688 (BRIT, W, WU). San Martin:

64 Sipa 18(1)

Lamas, on old trail from San Antonio de Cumbasa, S of Shapajilla, upper slopes of ee
sco, 06° 22'S, 76° 23' W, 600-800 m, 5 Oct 1986 (bud), S. Knapp et al. 8514, (fr), :
Knapp et al. 8517 (MO. US, USM).

Cybianthus spicatus is a polymorphic ochlospecies, sensu White (1962),
Prance (1972) and Pipoly (1983a), with many semi-isolated populations
throughout the Amazon and eastern Guayana Floristic Province (Maguire
1979). These localized populations have produced several seemingly dis-
tinct ecotypes, resulting in overdescription. Collections of Cybianthus spicatus
from Peru match the type of Conomorpha gracilis A. C. Smith, described from
Guyana, in every detail. Fieldwork conducted in Guyana, Venezuela, Bra-
zil and Peru since my earlier treatment (Pipoly 1983a), where I discuss the
four ecotypes present within the species, and the synonymy rationale, has
confirmed the concept that Cybianthus spicatus is a polymorphic ochlospecies.

Steyermark (1981) discussed the fact that Richard Schomburgk'’s collec-
tions attributed to Guyana are now known to be from Venezuela.

14. Cybianthus fulvopulverulentus (Mez) G. Se subsp. magnoliifolius
(Mez) Pipoly (Fig. 3A), Brittonia 35:72. 1983. Conomorpha magnoliifolia
Mez in Engl., Pllanzenr. IV. 236(Heft 9):258. 1902. Cybianthus saa (Mez)
G. Agostini, Acta Biol. Venez. 10:146. 1980. Typz: SURINA ichout cle.
(scam. fl), H. Kegel 244 (LECToTYPE by Pipaly 1983a: i

Conomorpha rigida Mez, Repert. Spec. Nov. Regni Veg. 16:420. 1920. syn. nov. Type:
L. AMAZONAS: “Hylaea,” without date (fr), E. U/e 8722 (noLorype: B-destr., F
Neg. 4831). rae searches of herbaria housing significant Ule collections, no oe
eae of the type have been located. Therefore, I select a neotype herewith: Type:

GUYANA [BRITISH GUIANA]. Orealla Savanna, Corentyne River, Sep 1879 (fr),
E. Imthurn B/9 (Neoryee here designated: K; lsONEOTYPE: BRG). The neotype was annotated
by Mez in 1901, but was not mentioned in the protologue.

>
~ rs

Shrub or tree to 10 m tall. Branchlets mostly thick, (3—)5.2—12 mm diam.,
glabrate or densely stellate and dendroid ferrugineous tomentose and glan-
dular-granulose, at times glabrescent. Leaves alternate, at times clustered
and approaching pseudoverticillate; blades coriaceous, elliptic, oblanceolate,
ovate or obovate, 7.2—21.7 cm long, (2.2—)3.6—9.5(—17.1) cm wide, apically
obtuse, acute, rounded or emarginate, basally obtuse to acute (rarely acuminate),
sparsely pitted with superimposed glandular lepidote scales and glabrous
or glandular-granulose, at times sparsely ferrugineous tomentose above and
below, glabrescent, epunctate, the margin entire and revolute; petioles marginate,
(1.3—)1.6—4.2(-4.6) cm long, ferrugineous tomentose, and glandular-granulose,
glabrescent. Staminate inflorescence: a simple, tortuous raceme, rarely lax, (3.6-)
4.5—15 cm long, the rachis maroon, opaque, glandular-granulose or with a
few scattered dendroid ferrugineous trichomes, glabrescent; floral bracts
chartaceous, linear-lanceolate, 0.8—3.6 mm long, ca. 0.1 —0.2 mm wide, densely
ferrugineous tomentose, caducous; pedicels cylindrical, (0.2—)2.9—3.6 mm

Prpoty, Cybianthus in Ecuador and Peru 65

long, densely glandular-granulose, glabrescent. Staminate flowers 4- or 5-merous,
chartaceous, white, cream or yellow; calyx deeply cotyliform, 0.9—2.6 mm
long, densely glandular-granulose then glabrescent, the tube 0.3—0.7 mm
long, unequally divided, the lobes chartaceous, ovate to ovate-triangular,
0.6-1.9 mm long, 1.1—2.4 mm wide, obtuse to acute or acuminate apically,
prominently orange punctate, densely ferrugineous tomentose and ferrugi-
neous glandular-granulose or partially glabrescent or ferrugineous glandu-
lar-granulose and them sometimes totally glabrescent, the margins extremely
undulate, entire, densely glandular-ciliate; corolla campanulate, (2.8—)3.3—
4.7(—5.5) mm long, the tube (0.6-)0.9-1.4(-1.7) mm long, the lobes ovate,
(2—)2.2—-3.1(—4) mm long, (1.2—)1.5—2.4(-2.8) mm wide, the lobes highly
reflexed at anthesis, obtuse to acute apically, conspicously orange punctate
and punctate-lineate, at times scattered ferrugineous trichomes before an-
thesis without, the margin entire; stamens (2.4—)2.8—3.4 (—3.8) mm long,
the staminal tube 0.8—1.1 mm long, the apically free portions of the fila-
ments 0.6—0.9 mm long, the anthers elongate-triangular, (0.8—)1—1.4 mm
long, apically acute, basally cordate, dorsifixed ca. 1/4 from base, distally
recurved or rarely and aberrantly erect; pistillode (1.6—)1.9—2.5 mm long,
the ovary 0.7—1.1 mm long, 0.7—1.2(-1.4) mm diam., densely translucent
glandular-lepidote, the style elongate, curved, 1-1.6 mm long, the stigma
0.1-0.2 mm long. Pistillate inflorescence as in staminate but erect (3—)4—11.4
cm long, the rachis green then red; floral bracts 0.7—2.9 mm long, ca. 0.1—
0.2 mm wide; pedicels cylindrical, (0.2—)0.6—1.5(—2) mm long. Pusti/late
flowers as in staminate but dull yellow to brown; calyx 1.2—2.6 mm long,
the tube 0.4—0.7(—1) mm long, the lobes 0.7—2 mm long, 1.3—2.4 mm wide;
corolla (2.7-)3.2—3.5(—4) mm long, the tube 0.6—-1.2 mm long, the lobes
2—2.3(—2.6) mm long, 1.1—1.5 mm wide; staminodes as in stamens but vee
2.9 mm long, the staminodial tube 0.7—0.9 mm long, the apically free portions
of the filaments 0.7 mm long, the antherodes triangular, 0.6—0.9 mm long;
pistil obnapiform, 1.9—2.5(—3) mm long, the ovary (0.7—)0.9-1 .6 mm long,
(0.7—)1.3-1.7(—2.2) mm diam., densely translucent glandular-lepidote, the
style thick, (0.4—)0.6—0.9(-1.3) mm long, the stigma pseudocapitate, very
minutely 4-lobed, up to 0.3 mm long. Fruit globose, 3—9 mm long and in
diam., green, then red, then black, inconspicuously pellucid punctate.

Distribution.—Venezuela, Guyana, Surinam, French Guiana, Brazil (Amazonas,
Roraima, Parad, Mato Grosso) and reported here for the first time from Peru
and Bolivia 50O—500(—850) m.

Ecology and conservation status —Cybianthus fe
is extremely common in wet tepuf savannas, but is rare in Peru it in varillal
htimedo (wet sandy, non-inundating) habitats. It is anticipated, but has not
yet been collected, in Ecuador.

Etymology.—The epithet, ‘fulvopulverulentus’ refers to the rusty tomen-

Ped) | en 1 yee ys
Lf

66 Sipa 18(1)

tum of the species, forming a powdery like vestiture on the adaxial leaf surface,
caducous as the leaf matures. The subspecific epithet refers to the shape and
aes adaxial leaf surface, giving the general impression of a Magnolia leaf.

Specimens ined. PERU. Loreto: Prov. Maynas, Iquitos, Nina rumi-Rio Nanay, 03°
48! S, 73° 25' W, 122 m, 5 Mar 1987 (bud), R. Vasquez et al. 8905 (AMAZ, MO, US,
USM). BOLIVIA. Santa Cruz: Prov. Velasco, Parque ee Noel Kempff Meeade,
Campamento Huanchaca I, 3.35 km from Rio Pauserna, 13° 56! OL" S, 60° 49' 30" W,
600 m, 22 May 1994 (fr), L. Arroyo et al. 738 (BRIT, MO, Sa 26 Jul 1995 (pise. fl), 7
fee et al. 7528 (BRIT, MO, USZ), (stam. fl), T; Killeen et al. 7530 (BRIT, MO, USZ),

3 Nov 1995 (fr), A. Rodriguez & J. Surubé 669 (BRIT, MO, USZ).

(eee cited here are the first known from Peru and Bolivia. Those
collections represent remarkable disjunctions, the closest population known
heretofore occurring approximately one degree north of the Rio Negro in
southern Venezuela near the Brazilian frontier, or halfway across Brazil to
the east, on Serra do Cachimbo, state of Mato Grosso. To date, no speci-
mens are known from Ecuador.

In my previous treatment (Pipoly 1983a), I included Conomorpha rigida
in the synonymy of Cybianthus fulvopulverulentus (Mez) G. Agostini subsp.
fulvopulverulentus. The type at Berlin had been destroyed and I synonymized
it based on the description and photo. However, the discovery of two du-
plicates of the Imthurn gathering annotated by Mez permitted neotypification
of the name with the Kew specimen and its inclusion here. Populations
represented by this collection differ from the type of subspecies magnoliifolius
only in their diminutive stature and vegetative parts.

IV. ee ame subgenus Comomyrsine (Hook. f.) G. Agostini, Acta Biol.
z. 10: a 1980. Comomyrsine Hook. f. in Benth. & Hook., Gen. Pl. 2:643. 1876;

ees A. DC. subgenus ae ares f.) Mez im Engl., Phanvens IV. 236(Heft
9):290. ar So in Engl. & Pflanzenfam. IV, 1:92. 1897; J.F. Macbr., Field
Mus. Nat. Hist., Bot. Ser. 136, 1): i 1959. Type Species: Cybianthus a laok.

f.) G. da Geeonce: by D’Arcy, Ann. Missouri Bot. Gard. 60:445. 1973.)

Terrestrial, erect, monoaxial swbshribs or treelets to 2(—S) m tall. Roots positively
geotropic. Bark mostly brown, smooth, longitudinally fissured or transversely
checked. Trunk distinguishable, leptocaulous, monoaxial, following mor-
phogenetic dynamics of Corner’s Architectural Model (Hallé et al. 1978),
growth rythmic. Svems terete distally, sparingly to densely glandular-papil-
late, at times with rufous hydropotes or orange glandular lepidote scales,
without lenticels. Cataphylls alternate or pseudoverticillate, alternating with
pseudoverticels of leaves, or apparently axillary to them, linear-subulate to
acicular, rigid to membranous, keeled or flat, prominently punctate or punctate-
lineate, glabrous, glandular-papillate, bearing hydropotes, or orange lepi-
dote scales. Leaves pseudoverticillate or alternate, apically acute to rounded,
often mucronulate, basally symmetric or asymmetric, acute, obtuse, or ta-

PipoLy, Cybianthus in Ecuador and Peru 67

pering, rarely auriculate, the venation camptodromous, to brochidodromous,
apically acute or attenute to rarely rounded, mucronulate or not, basally
acute, cuneate, or rarely obtuse, at times subauriculate, symmetric or asymmetric,
black or pellucid punctate, bearing hydropoten, at times with orange glan-
dular lepidote scales or glandular papillae, the margin opaque or scarious,
entire, subentire, undulate and sparse dentate or pectinate-dentate, flat or
inrolled, the teeth vascularized or not; petioles pulvinate (basally swollen),
canaliculate, except rarely marginate distally. Inflorescence pinnately to tripinnately
paniculate, (rarely reduced to a pseudoraceme), pyramidal to columnar, the
branches spicate or racemose (rarely glomerulate); inflorescence bract lin-
ear-subulate, indistinguishable from the cataphylls; branch bracts membranous,
linear, subulate; floral bracts linear-lanceolate, subulate, subtending or rarely
on the pedicel, variously punctate, glandular-papillate or glabrous; pedicels
terete, thin, or obsolete. Flowers unisexual or bisexual, 4—5(—6)-merous,
homomerous or heteromerous; calyx cotyliform, at times unequally divided,
the lobes valvate, spreading, linear-subulate or rarely subdeltate, apically
acute, attenuate or rarely premorse, pellucid to black punctate, the margin
entire or erose-serrulate, glabrous or glandular-ciliate; corolla rotate to subrotate,
the lobes valvate, linear-subulate, apically acute, rounded or attenuate, es-
sentially glabrous without, densely glandular-granulose throughout within,
pellucid or black punctate, the margin irregular, entire; stamens and staminodes
similar, the staminodes reduced in size, partially connate to form a con-
spicuous or inconspicuous tube, at times bearing lobes alternating with apically
free filaments, the tube adnate to the corolla, at times so as to mimic epipetalous
stamens, the apically free portions of the filaments terete or flat, glabrous
or glandular-granulose, the anthers dorsifixed, as wide or wider than long,
apically rounded, obtuse or emarginate, basally cordate to deeply cordate,
dehiscent by wide longitudinal slites; pistillode lageniform, clavate or conic,
hollow, or entirely absent; pistil obpyriform, subglobose, obturbinate or clavate,
the ovary terete, sparsely to densely translucent glandular lepidote, glan-
dular-papillate or glabrous, the placenta umbonate, 2—3(—4)-ovulate, the
ovules immersed in the placenta ca. 1/2 their length, the style short or ves-
tigial, the stigma large, capitate, the margin lacinate, the lacinae with large
lobes, the stigma early caducous. Fruit globose to depressed-globose, the
exocarp sometimes fleshy, the mesocarp and endocarp stony, black or pellu-
cid punctate, one-seeded, the testa corrugate, the embryo cylindrical, transverse.

Distribution. —From the Darién of Panama southward through the Andes
of Colombia, Ecuador, Peru and Bolivia to Rond6énia, Brazil, at elevations
of sea level—2,200 m elevation.

Ecology.—Members of Cybianthus subgenus Comomyrsine occupy the for-
est floor, growing in sheltered areas under cliffs, and in other shady areas.
The monoaxial stems are weak and break easily when trampled. They are

68 Sipa 18(1)

extremely sensitive to soil compaction and occur mostly in areas where leaf
litter accumulates.

Pipoly (1987) demonstrated that Cybianthus subgenus Comomyrsine is most
closely related to subgenus Triadopbora by the monoaxial habit. The cataphylls
of Comomyrsine without petiolar structures, as opposed to the petiolate
pseudocataphylls found in subgenus Triadophora, serve to further distinguis|
Comomyrsine. Likewise, the caducous, oversized stigma with lacinate lobes
is unique to subgenus Comomyrsine. The subgenus contains 8 species, of which

7 occur in Ecuador and Peru.
KEY TO SPECIES OF CYBIANTHUS SUBGENUS COMOMYRSINE

oo 5.7—29 cm long; stems less than 1 cm diameter.

2. Stems apices bearing orange hydropotes; leaf blades linear-lanceolate, 1.6—

4.0 cm wide, the margin repand to undulate; petioles 0.6—1.0 cm i
inflorescence pseudoracemose, the flowers glomerulate. ........... 15. C. verticilloides
2. Stem apices bearing rufous glandular-papillate or with rufous hydropotes;

leaf blades ovate, obovate, oblanceolate, elliptic, or oblong, (4.0-),

22(—30) cm wide, the margin regular, entire or minutely straight-serru-
ate; petioles (1.0—)1.5—5(—21) cm long; inflorescence pyramidal-bipin-
nate paniculate, or a columnar thyrsoid panicle, never a pseudoraceme

the flowers spicate, corymbose, or rarely cymose.

. Stem apices sparsely glandular-papillate, bearing rufous hydropotes; leaf

ee broadly rounded, the mareins scarious; petioles progressively longer

n long; inflorescence paniculate,
ee mm long; sepals deltate; pet-

acropetally along the s

the flowers corymbose; pedicels 1
als cucullate apically, the margin glandular-granulose; filaments
6. C. croatii

glabrous.
3. Stem apices densely glandular papillate, without hydropotes; leaf bases
acute, to acute with a rounded base, the margins opaque; petioles subequal
along stem, slightly shorter acropetally, 1.6—2.0 cm long; inflorescence
pinnately paniculate, the flowers spicate or racemose; pedicels absent
abrous; filaments gland ae

sepals ovate-lanceolate; petals flat, the margin g!
7.C, humilis

granulose
1. Leaves (16—)31—105 cm long; stems 1—5 cm diameter.
4. Leaves oblong to elliptic, apically rounded or rarely acute, not mucronulate,
basally abruptly acuce to broad| 2 rounded; petioles ((2.5—)5 cm long; flowers
homomerous, 4— or
Shrub or tree to 5 m tall; in seine a pyramidal panicle, t
spicate, the es sessile to subsessile; leaf base symmetric; —

he branches

.C. sprucei

(2.5—) 3.5-5.0 cm long
5. Semi-woody nae to 1(—3.5) m tall; inflorescence a columnar, iyi
panicle, the branches racemose, the flowers on pedicels 2.3 m
long; leaf base asymmetric; cataphylls 0.6-2.5 cm long. wo... 19. C. simplex
apically acute, mucronulate, basally gradually tapered
—2 cm long; flowers het-

oo iees Gisneealak

on the setiole. often sub auriculate; petioles 1.5
eromerous, the calyx 5—6-lobed, the corolla 4—5-lobed,
6. Leaves ee ae 31-55 cm long, the margin flat, entire; cataphylls

Pipoty, Cybianthus in Ecuador and Peru 69

1.5—4.0(-4.5) cm long; calyx lobes delrate, the margin glabrous; co-
rolla carnose 20. C. kayapit

6.1 riaceous, longer than 5S—1 25cm, the margin inrolled, densely
and eens serrulate; cataphylls 4.0-6.5 cm long; calyx lobes linear-
subulate, the margin densely glandular-granulose; corolla membrana-
ceous. 1. C. anthuriophyllus

1: Laer verticilloides (Cuatrec. )G. ake (Fig. 4B), Acta Biol.
Vene 0:165. 1980. \ ., Revista Acad. Colomb. Ci.
Exact. ae 1951. TyeE COLOMBIA. ie DEL Cauca: Rio Cali riverbed,
between Quebrada de Juntas and El Recreo, 2,070-2,260 m, 7 Jul 1946 (stam. fl), J.
Cuatrecasas 21981 (HOLOTYPE: F; IsoTypE: COL).

Shrub to 1 m tall. Stem terete, 5-8 mm diam., sparingly rufous glandu-
lar-papillate and bearing orange hydropotes, early glabrescent, the bark
horizontally checked. Cataphyl/s chartaceous, subulate, (1.2—)1.5—2.1 mm
long, 0.9-2.1 mm wide, flat, apically long-attenuate, basally somewhat decurrent,
sparingly glandular-papillate and orange lepidote above and below, black
punctate. Leaves pseudoverticillate; blades chartaceous, linear-lanceolate, 1 2—
20 cm long, 1.6—4.0 cm wide, apically long-attenuate, basally acute, grey-
ish-green above and pallid green below when fresh (teste Cuatrec.), spar-
ingly rufous glandular-papillate and orange lepidote above and below at
first, persistent only below, inconspicuously pellucid-punctate, midrib planar
above, prominent below, the secondary veins 8—12 pairs, prominent below,
the margin inrolled except revolute basally, undulate to a vascularized blunt
tooth at nerve end, translucent throughout its length but not scarious; petioles
canalicualte, 0.6—1.0 cm long, 0.5—1.0 mm diam., glabrous, swollen ba-
sally at point of attachment. Staminate Inflorescence: a pseudoraceme, 10-12
cm long; peduncle ca. 1 cm long, the rachis glandular-papillate, the flowers
glomerulate; inflorescence branch bracts membranous, subulate, 5.5—7.0
mm long, 1.2-2.0 mm wide, minutely glandular-papillose publerulent,
glabrescent, prominently black lineate-punctate, the margin entire; floral
bracts similar to branch bracts except 0.9—2.0 mm long, 0.2—0.5 mm wide;
pedicels obsolete to cylindrical to 1.3 mm long, prominently black punc-
tate. Staminate flowers 5—6-merous, lilac; calyx membranaceous, subcotyliform,
1.5—2.3 mm long, unequally divided, the tube to 0.2 mm long, the lobes
linear-lanceolate, subulate, 1.3—2.0 mm long, 0.4—0.7 mm wide, apically
long-attenuate, flat, epunctate, glabrous, the margin irregular, entire, gla-
brous; corolla rotate, chartaceous, 2.5—4.0 mm long,the tube opaque, 0.8—
1.5 mm long, the lobes hyaline, ovate, 2.1-3.0 mm long, 1.4—1.9 mm wide,
reflexed 180°, apically acute to rounded, moderately glandular-granulose
throughout within except densely so along margin, glabrous without, epunctate,
the margin irregular, entire; stamens 3.0—-3.5 mm long, the tube 0.8—1.5
mm long,epunctate, elobate, glabrous, the apically free portions of the fila-

70 Sipa 18(1)

ments thick (base of filament as wide as anther), terete, glabrous, 1.5—2
mm long, ventrally reflexed apically, the anthers subglobose, wider than
long, 0.3-0.5 mm long and wide, apically rounded to obtuse, basally barely
cordulate, the connective prominately black punctate dorsally; pistillode
ovoid, ca. 0.9-1.3 mm long, 0.6-0.8 mm diam. Pistillate inflorescence resem-
bling staminate, but 4.5—6.5 cm long, the glomerules mostly reduced or a
single flower; branch bracts and floral bracts identical, membranous, sublate,
2.8—4.0 mm long, 0.7—1.0 mm wide, sparsely glandular-papillate, promi-
nently black punctate, the margin erose; pedicels (1.5—)1.8—3.2 mm long.
Pistillate flowers as in staminate but white; calyx chartaceous, cotyliform,
2.2—2.7 mm long, hyaline, the tube 0.3—0.4 mm long, the lobes 1.9—2.3
mm long, 0.7—1.1 mm wide, apically attenuate, at times rufous-papillate
apically; corolla rotate, 2.5—3.2 mm long, the tube 0.9-1.0 mm long, the
lobes widely ovate, 1.6—2.2 mm long,1.2—1.5 mm wide, apically rounded
to acutish, glandular-granulose within and along margin, the margin regular;
staminodes similar to stamens but 1.6—1.9 mm long, the tube 0.9—1.0 mm
long, the apically free portions of the filaments terete, 0.7—0.9 mm long,
the antherodes globose, often malformed, 0.2—0.3 mm long, 0.3—0.4 mm
wide, apically obtuse, basally obtuse to rounded or cordulate, the connec-
tive prominently black punctate ventrally; pistil obpyriform, 2.1—2.5 mm
long, 1.2—1.6 mm diam, the ovary 1.6—1.8 mm long, translucent-lepidote,
glandular-papillate, prominently black punctate, style 0.5—0.7 mm long,
the stigma capitate, 4—5-lobed, each lobe lacinate, to 0.6 mm long, the placenta
deeply cupuliform, 2 ovulate. Frit globose, 7—7.5 mm long and diam.,
prominently black punctate, exocarp thin.

Distribution. —Previously considered endemic to the Western Cordillera
of the Andes, on the cliffs (Farallones) of Cali, in the Department of Valle,
Colombia, at 1,900—2,600 m, but reported for ie first time here from Napo,
Ecuador, at 200 m elevation.

Ecology and conservation status. —Cybianthus verticilloides is a rare species in
lowland and premontane forest, growing near watercourses. Given that it
has well-known uses, C. vertici/loides might be suitable for cultivation. However,
at present nothing is known regarding its natural population dynamics. Given
the rapid urban development around Cali, and its apparent rarity in Ecua-
dor, this species should be considered endangered.

Etymology.—The specific epithet refers to the strikingly pseudoverticillate
phyllotaxis.

Local names and uses.—Colombia: “verticilado” (Spanish); Ecuador:
“Carnerupachapanga,” “Yanacarneru” (Quichua), “Carnero Negro” (Spanish).
Used to get rid of small fish in the urinary tract that cause bleeding and pain.

cimens examined. COLOMBIA. Valle del Cauca: Rio Cali riverbed, above R fo Pichindé,
El een 2,640 m, 25 Jul 1946 (fr), J. Cuatrecasas 21721 (COL, F); Pefias Blancas, 2,200

Prpoty, Cybianthus in Ecuador and Peru gia

m, 9 May 1940 (stam. fl), A. Figueroa 875 (COL,US); Rio Tulud, 1,200 m, without date
(pise. fl), J. Dugue-Jaramillo 4022 (COL); El Silencio, Yanaconas, 1,900—2,200 m, 28 Feb

1939 (pist. fl), E. Killip & H. Garcia 33802 (COL, US); Farallones de Cali, 1,800 m, Oct
1883 (pist. fl), E Lehmann 3027 (K, US); km 18-20, Cali-Buenaventura Hwy, entering
near Finca Zingara, summit of the Cricilen Occidental, 1,500-2,000 m, 28 Feb 1988
(fr), H. van der Werff G I. Cabrera 15786 (COL, MO, VALLE). ECUADOR. Napo: Rio
Napo, S bank a few km below Itaya, 00° 28'S, 76° 3 33' W, 200 m, 20 Aug 1982 (ster.) H.
Balslev & Santos Dea 2850 (QCA)

Cybianthus verticilloides appears to be most closely related to C. goudotianus,
by the synapomorphic vascularized leaf teeth. However, C. verticilloides is
defined by the autapomorphic horizontally checked bark, the orange hydropotes
of the branchlets, and the linear-lanceolate leaf shape.

16. Cybianthus croatii Pipoly, sp. nov. (Fig. 4A, 16). Tyre: Ee UADOR. Pastaza:
between sbi z de Agosto and Arajuno, 18 km NE of main Puyo-Macas
Rd., 8.2 km NE of Diez de Agosto, 01° 27'S, 77° 51' W, 970 m, 4 May 1984 (stam.

fl), i Ge 59009 ees PE: MQ).

Quoad habitum deminutum petiolos brevistipitatos et laminas chartaceas, C. humilem
valde cognatum, sed ab ea basibus laminaribus obtusis vel rotundatis (non acutis) inflorescentiis
paniculatis cum ramulis floriferis corymbosis (nec tripinnati- paniculatis cum ramulis floriferis
spicatis vel racemosis) petalis ad apicem cucullatis (nec planis) ad marginem glandular-
papillosis (nec glabris) filamentis glabris (nec glandulari-granulosis) antherarum connectivis
manifeste punctatis (nec epunctatis) prompte cognoscitur.

Subshrub to ca. 15 cm tall. Stem terete, ca. 4 mm diam., bearing rufous
hydropotes and sparingly glandular-papillate. L Iternate; blades chartaceous,
ovate to elliptic, 5.7-14 cm long, 4.0—7.5 cm wide, apically acute, the tip
mucronulate, basally obtuse to rounded slightly decurrent on the petiole,
bearing rufous hydropoten above and below, midrib slightly impressed distally,
slightly raised proximally above, prominent below, the secondary veins 4—
9 pairs, barely visible above, prominent below, without collecting vein, the
margin scarious, subentire or entire; petiole deeply canaliculate, decurrent
on stem, at times appearing to form a small sheath, (1.5—)2.0—4.5 cm long,
().2—-0.3 cm diam., increasing in length acropetally along stem. Cataphylls
membranaceous, alternate, subulate, 6-12 mm long, 0.6-1.2 mm wide,
located just below center of internode, psuedoverticillate, apically acicular,
keeled, conspicuously black punctate-lineate, bearing hydropoten and glandular
papillae. Inflorescence bract subulate, 3.9-4.0 mm long, 2.0-3.0 mm wide.
Staminate inflorescence: supraaxillary, paniculate, 1.2—3.5 cm long, appearing
succulent, the branches racemose, the rachis densely glandular-papillate;
peduncle 0.5—1.0 cm long; inflorescence branch bracts linear-lanceolate, 1.8—
2.2 mm long, 0.4—0.6 mm wide, conspicuously punctate, glandular-papil-
late; floral bracts subtending and equal to the pi 1.0-2. > mm long.
Staminate flowers homomerous, 4-merous, green; cal tyli

1.2-1.5 mm long, more or less equally divided, hyaline, the tube 0.2—-0. 3

72 Sipa 18(1)

Fic. 16. Cybianthus croatii Pipoly. A. Habit, showing stem with small, acicular cataphylls,
acropetally longer petioles, and supraaxillary inflorescences. B. Staminate flower bud, showing
sparsely seen papier’, oe coke C. Open staminate corolla, aowine cucul-

late lobe apic conic pistillode. D. Open pistillate corolla, showing
oblate ee. subglobose pistil. A-C, drawn from holotype. D, drawn from L. Adhert
de Escobar 3744. Figure drawn by Peggy Duke.

Prpo.y, Cybianthus in Ecuador and Peru 73

mm long, the lobes deltate, 1.0—1.2 mm long and wide, apically acute to
somewhat acuminate, prominately black punctate, sparsely glandular-papillate
without, the margin irregular, entire, glandular-papillate at first, glabres-
cent; corolla subrotate, carnose, 2.4—2.7 mm long, the tube 0.7—0.8 mm
long, translucent, the lobes opaque, oblong, 1.7—2.1 mm long, 0.9-1.0 mm
wide, apically rounded to obtuse, cucullate, sparingly glandular-papillate
without, densely glandular-granulose within, prominently black punctate
without, especially at apex, the margin entire, glandular-granulose; stamens
2.5—2.6 mm long, the tube conspcuous, carnose, ().7—0.8 mm long, subtruncate,
the apically free portions of the filaments terete, 0.9—1.0 mm long, epunctate,
glabrous, erect except slightly recurved ventrally at point of attachment to
anther, the anthers suborbicular, (.7—0.8 mm long and wide, apically rounded,
deeply cordate basally, the connective epunctate ventrally, prominently black
punctate dorsally; pistillode conic, 1.3 mm long, 0.7 mm wide, densely
translucent-lepidote, conspicuously black punctate, hollow. Pzstz//ate inflorescence:
as staminate, but 2.5—3.5 cm long; pedicels 0.6—1.1 mm long. Paéstillate
flowers as in staminate but calyx 1.3—1.5 mm long, equally divided, the tube
0.2—0.3 mm long, the lobes triangular, 0.9—1.2 mm long, 0.5—0.7 mm wide,
apically acuminate, sparsely glandular-papillate along margin without; corolla
subcampanulate, the lobes 1.3—1.5 mm long, the tube to 0.2 mm long, the
lobes suborbicular, 1.1—-1.3 mm long, 1.0—1.1 mm wide, apically rounded
to obtuse, minutely glandular-granulose along margin without and throughout;
staminodes as in stamens but 0.9—1.0 mm long, the tube, ca. 0.2 mm long,
the apically free portions of the filaments 0.2—0.3 mm long, the antherodes
oblate, 0.4—0.5 mm long, 0.5—0.6 mm wide, apically truncate, basally obtuse,
the connective prominently black punctate ventrally; pistil subglobosel, 0.6—
0.7 mm long and diam., densely translucent-lepidote, conspicuously black
punctate, the placenta deeply cupuliform, bearing 2 ovules, the style barely
discernible, to 0.1 mm long, stignma subcapitate, the margin lacinate, early
caducous. Fruit unknown.

Distribution.—Known only from the type (Ecuador) and from Antioquia,
Colombia, at 970—2,440 m elevation..

Ecology and conservation status. —Cybianthus croatii occurs in premontane
and montane wet forest, where it is locally common in protected areas near
forest margins. Because of its restricted habitat, it is presumed to be threatened.

Etymology.—It is with great pleasure that I dedicate this species to Tho-
mas B. Croat, pre-eminent authority on the systematics of Neotropical Araceae,
indefatigable collector, gentleman, scholar, and the P. A. Schulze Curator
of Botany at the Missouri Botanical Garden. During the tropical botany
course I took from him in Costa Rica in 1977, I was first shown and be-
came intrigued with the systematics and populaton biology of the Myrsinaceae.

74 Sipa 18(1)

ParatyPe: COLOMBIA. Antioquia: Mpio. Caldas, above town, Finca La Zarza, 2,440

, 2 Feb 1984 (pist. Al), L. Albert de ty et al, 3744 (HUA, US

Cybianthus croatii is the smallest myrsinaceous shrub that exhibits Corner’s
Model (Hallé et al. 1978). It is interesting to note that with each successive
pseudowhorl of cataphylls, an inflorescence is produced, followed by a larger
leaf with a longer petiole. This species appears to be most closely related to
another diminutive plant, C. Avmilis, known only from the Department of
Antioquia, Colombia and adjacent Chocé, and from Ecuador. The synapomorphy
which defines the Cy a humilis-C. croati clade is the scarious leaf margin,
known elsewhere only in subgenus Grammadenia (Pipoly 1987, unpubl. data).
However, because that subgenus occurs on the other side of the generic cla-
dogram presented by Pipoly (1987), it is most parsimonius to hypothesize
that the margin has arisen independently in these distant lineages. The pistillode
is present in both of these species, and I have chosen to postulate that this
represents a reversal because it is the most parsimonius conclusion. Cybianthus
croatil is defined by the autapomorphic rufous stem hydropotes and the ac-
ropetally longer petioles. Despite the fact that the distribution of this spe-
cies entirely overlaps that of C. Awmulis, it appears that C. croatii is restricted
to montane and cloud forests, where C. Awmi/is is restricted to premontane
pluvial forests and subparamo thickets. This is yet another example of sis-
ter species in altitudinally adjacent habitats, already reported in Cybianthus
subgenus Laxiflorus (Pipoly 1983) C. subgenus Microconomorpha (Pipoly 1983b)
and C. subgenus Conomorpha (Pipoly 1992a). Whether this supports the concept
of speciation by peripheral isolation (parapatric) is unknown. Futher stud-
ies of the respective population biologies of the subgenus are needed.

17. Cybianthus humilis (Mez ) G. Agostini (Fig. 3F), Acta. Biol. Venez.
10:163. ehen oe Mez in Engl., Pflanzenr. IV. 236(Heft 9):291. 1902
Type: ECUADC hout oie 1896 (stam. fl), A. Sodiro 100/14 (HoLoTyPE:

-destr., F Neg pe LEC here designated: COLOMBIA. ANtTIoQuia: 2,650—

2,800 m, AGE 1880 ee a W. <albreyer 1534 UK). Although Mez (1902) did
not specifically mention the Sodiro collection as the type, he cited the Sodiro and
a ne Haye er ne in the Pioreiogye Blow aves the FP phorograph clearly shows

- that of Kalbreyer
does not, ene ene that che Bodine specimen formed the e principal pie upon
which the aoe was . ised. eae tunately, : :

have been located. 1 the ee sheet at K, and thus I meee ite
it as the lectotype.

oe hella Mez, Bull. Herb. Boissier, 2 ser 5:535. 1905. syn. nov. Cybianthus
x. Agostini, Acta Biol. Venez. 10:163. 1 1980. ae ECUADOR. Atacarzo.

oe 1902 (stam. fl), A. Sodiro 100/2 (HOLOTYPE: B-destr.; LECroryeE, here designated: P).
Subshrub to 25 cm tall. Stem terete, 3.5—4.0 mm diam., densely glandu-
lar-papillate, early glabrescent. Cataphy//s in alternating nodes with leaves,
membranaceous, subulate, 10-13 mm long, 1.7—2.6 mm wide, apically long-

Pipoty, Cybianthus in Ecuador and Peru Te

attenuate, keeled, midrib prominent below, prominently black punctate and
lineate-punctate, glandular-papillate, glabrescent, the margin opaque, flat,
entire. Leaves subopposite; blades chartaceous, elliptic to narrowly elliptic,
15—20(—27) mm long, (4.0—)7.0-8.5 cm wide, apically acute to subacuminate,
terminating in an inconspicuous mucro, basally acute, bearing hydropoten
above and below, inconspicuously pellucid punctate, midrib somewhat impressed
above, prominent below, the secondary veins 6—11 pairs, slightly impressed
abive or not, prominently raised below, not united by a submarginal col-
lecting vein, the margin entire, opaque, subrevolute; petioles canaliculate,
thin, 1.0—2.0 cm long, 2.0—2.5 mm diam., glabrous, not decurrent on the
stem. Inflorescence bracts similar to cataphylls, but 1.2-1.3 mm long, 1.5—
2.2 mm wide. Staminate inflorescence: a bipinnate panicle, 3.5—15(-19) cm
long, the rachis densely glandular-papillate, the branches spicate or rarely
racemose, appearing subglomerulate apically; inflorescence branch bracts
membranaceous, subulate, 3—4.5 mm long, 0.3—0.5 mm wide, densely and
prominently black punctate-lineate, sparingly papillate; pedicels essentially
obsolete, to 0.3(—2) mm long; floral bract membranaceous, ovate, asym-
metric, 2.3-2.8 mm long, 0.9-1.2 mm wide, apically abruptly acuminate,
medially keeled, epunctate, the margin erose, stipitate glandular-papillate.
Staminate flowers 4—S-merous; calyx membranaceous, cotyliform, 1.5—2.0 mm
long, tube 0.2-0.3 mm long, the lobes ovate to lanceolate, 1.3—1.7 mm
long, 0.5—1.3 mm wide, unequally divided, apically acuminate to acute and
often erose or premorse, glandular papillose-puberulent without, the mar-
gin erose, glabrous; corolla chartaceous, appearing subrotate, 3.7—4.5 mm
long, the tube 1.1—1.5 mm long, the lobes linear-lanceolate to oblong, 2.4—
3.2 mm long, 1.0—1.5 mm wide, often unequal, apically long-attenuate to
obtuse, moderately rufous gladular-granulose throughout within, sparsely
glandular-papillate without, hyaline, the margin entire, glabrous; stamens
3.0—3.9 mm long, the tube 1.1—1.5 mm long, coriaceous, conspicuous, sparsely
glandular-grandulose, elobate, the apically free portions of the filaments basally
as wide or wider than anther, then tapering apically, 1.9—2.4 mm long, terete,
glandular-granulose, the anthers oblate, 0.4—0.5 mm long, 0.6—0.7 mm wide,
apically truncate, basally subcordate, the connective epunctate; pistillode
glabrous lagenform, 1.4-1.6 mm long, 0.8—1.0 mm wide, the stigma punctiform.
Pistillate inflorescence: as in the staminate but 7-10 cm long, 4-6 cm wide;
inflorescence branch bracts 2.5—3.5 mm long, 0.2—0.3 mm wide; peduncle
1-2.5 cm long; floral bracts 1-1.5 mm long; pedicels obsolete. Pzstz/late
flowers unknown; fruiting calyx as in staminate but 1.3—1.5 mm long, the
tube ca. 0.2—0.3 mm long, the lobes 1.1—1.2 mm long, 0.2—0.3 mm wide.
Fruit globose, reddish-purple when fresh, 4-6 mm long and diam. when
dried, the exocarp thin, conspicuously pellucid punctate.

76 Sipa 18(1)

Distribution.—Known only from Colombia and Ecuador, at 1,450—2,700
m elevation.

Ecology and conservation status. —Cybianthus humilis grows in deep shade
among rocks at the margins of premontane pluvial forests. Populations I
observed in Colombia grew only in undisturbed areas with deep shade, leaf
litter and organic humus. Because of the apparently restricted habitat, this
species should be considered threatened

Etymology.—The specific epithet refers to the low habit of the plant.

Local names and uses, —Colombia (Chocé6): “Hierba del palo grande.” Ground
to make crude syrups; syrup applied externally to cure cuts, taken inter-
nally to cure internal infections, clean the stomach and against chills. Given
that it is a shade species, the common name probably refers to its frequency
in shaded areas near large trees.

Specimens examined. COLOMBIA. Antioquia: Mpio. Urrao, Parque Nacional Las Orquideas,
Vereda Calles, permanent inventory for premontane pluvial forest, right bank of Rio Calles,
06° 32' N, 76° 19' W, 1,450 m, 27 Nov 1993 (fr), J. ae et al, 17186 (COL, JAUM,
MO), 1,450-1,500 m, 29 Nov 1993 (fr), J. Pipoly et al. 17361 (COL, JAUM, MO), with-
out eee and date, (stam. fl), F Lehmann s.n. (FE, K). Choco: a de Quibd6, Corregimiento
San Francisco Ich6é, Quebrada Caledonia along Caledonia Rd., 9 Apr 1987 (ster.), F. Garcta
G - Echavarria 259-A (COL, CHOCO, MO). Narifio: Mpio. jee Corregimiento
Ortiz y Zamora, Vereda El Barro, Reserva Natural Rio Nambjf, c re Altaquer,
ee occidentales de la Cordillera Occidental, 01° 18'N, 78° oa 7 i: ee 400 m, 3
Sep 1997 (ster.), J. Pipoly, A. Cogollo, et al. 21240 (BRIT, FMB, JAUM, PSO ‘ Ouindio:
Mpuo. De Salento, Estacién Navarco, Alto San Ignacio, 2,850 m, 23 Nov 1990 (stam. fl),
P. Franco et al. 3204 (COL, MO). Risaralda: Mpio. Sta. Rosa, Camino de Herradura etre
Termales y Paramo Sta. Rosa, Cordillera Central, vertiente Occidental, Hacienda El Margarital,
2,500 m, 18 Aug 1980 (fr), J. Ldrobo et al. ons (COL, MO). ECUADOR. Napo: Canté6n
El Chaco, Proyecto Hidroeléctrico Coca, Punto $T3, right bank of Rio Quijos, ca. 10 km
S of Reventador, 1,500 m, 3-5 re 1990 (Ee), W. is 5950 (MO, QCNE). Pichincha:
Canton Quito; Parroquia Calacalf, Reserva Geobotanica Pululahua, 00° O1' N, 78° 35' W,
1,800—2,000 m, 29 Jul 1989 ae fl), C. Cerén 7184 (MO, QCNA), Mind6, 26 Jun 1876
(stam. fl), E. André 3819 (K).

eon

Cybianthus humilis is most closely related to C. croatii, by virtue of its
synapomorphic scarious leaf margin. The autapomorphies that distinguish
C. humilis from all other species of the subgenus include the premorse apices
of the calyx lobes, the unequal corolla division, and the oblate anther shape.

aoa sprucei (Hook. f.)G. Agostini, Acta Biol. Venez. 10(2):164.
L980. Comomyrsine spruce eae f. iz Benth. et Hook., Gen. P1. 2:644. 1876. eae
spruce? (Hook. f£.) Mez in Engl., PHanzenr. IV. 236(Heft 9):291. 1902. Tyee:
DOR. [CHimporazo: W slopes of Volcan Chimborazo, 17 Jun 1860] a ae
Spruce 6144 (HOLOTYPE, K-2 sheets). Note: label on specimen does not indicate place
or date. Information was derived by comments regarding habit, etc. on the label which
matched data given in Spruce (1880).
Weigeltia panamensis Standl., Publ. Field Mus. Nat. Hist, Bor. Ser. 22:164. 1940. syn.
nov. Cybianthus panamensis (Standl.) G. Agostini, Acta Biol. Venez. 10:163. 1980. Type:

Pipoty, Cybianthus in Ecuador and Peru rei

PANAMA. Danrién: Cana, Cuasi Trail, Drto. Cheijana, 1000 m, 10 Mar 1940 (stam.
fl), M. E. Terry & R. A. Terry 1490 (HOLOTYPE: F-2 sheets; IsoTyPEs: A, MO).
Weigeltia purpurea Cuatrec., Revista Acad. Colomb. Ci. Exact. 8(31):326. 1951. Type:
COLO A. VALLE DEL CAUCA: Bahia de Buenaventura, Quebrada de San Joaquin,
0-10 m, 21 Feb 1946 (stam. f1), J. Cuatrecasas 19892 (HOLOTYPE: F; IsoTyPE: COL)
Shrub to tree to 5 m, flowering from less than 1 m. Stem terete, 1.3—2.5
cm diam. below uppermost leaves, swollen at nodes, semi-woody, glandu-
lar-papillate-puberulent, glabrescent. Cataphylls few, alternate, coriaceous,
subulate, (2.5—)3.0-5.0 cm long, 2-4 mm wide, strongly keeled, densely
glandular-papillose-puberulent, glabrescent, consipuously black punctate,
the margin opaque, regular, entire. Leaves pseudoverticillate; blades chartaceous,
widely oblong to elliptic, rarely widely obovate, (26—)31—75 cm long, (6.5—)
10—30 cm wide, apically rounded, obtuse or rarely acutish, not mucronulate,
basally abruptly subcuneate, asymetric, slightly decurrent on petiole, bear-
ing a few hydropoten above, moderately rufous glandular-papillate and with
a moderate number of hydropoten below, prominently red or black puctate,
the margin regular, opaque, entire; petioles stiff, (2.5—)5.0—-14(—23) cm long,
0.3-0.9 cm diam., slightly to moderately canaliculate, swollen basally, sparingly
glandular-papillate, glabrescent. Staminate Inflorescence: a pyramidal bipin-
nate thyrsoid panicle, 9-16.5(—19) cm long, 9—17(—22) cm wide, the branches
subspicate, the rachis densely glandular-papillose-puberulent; peduncle 0.5—
2.0 cm long; inflorescence branch bracts chartaceous, linear-lanceolate, 8.5—
12.9 mm long, 0.6-0.9 mm wide, apically narrowly acute, conspicuously
black punctate, densely glandular-papillate, gabrescent, flat, the margin opaque,
regular, entire; floral bracts membraneous, linear, 1.5—2.0(—2.5) mm long,
).5—0.6 mm wide, subulate, hyaline, densely glandular-papillose puberu-
lent, the margin entire, glandular-papillate; pedicels obsolete to 0.2 mm
long. Staminate flowers homomerous, 4-merous, coriaceous, subsessile, race-
mose, 4-5-merous, white in bud, green in anthesis, then crimson; calyx
coriaceous, subcotyliform, unequally divided, 0.9-1.1 mm long, the tube
0.1-0.2 mm long, the lobes linear-lanceolate, 0.7-0.9 mm long, 0.2—0.5
mm wide, apically subulate, keeled, brown punctate-lineate or punctate medially,
sparsely glandular-papillate, the margin irregular, subentire to erose, densely
glandular-ciliate; corolla subrotate, 2.0—2.9 mm long, the tube 0.2—0.3mm
long, the lobes narrowly ovate to lanceolate, (1.7—)1.9—2.3(—2.6) mm long,
0.9-1.0(—-1.5) mm wide, reflexed 135° from tube at anthesis, apically at-
tenuate, densely glandular-granulose throughout within and along margin
within and without, inconspicuously pellucid punctate, the margin entire,
somewhat irregular, densely glandular-granulose; stamens exserted to slightly
shorter than the corolla lobe, 1.6—2.2 mm long, the tube ca. 0.5 mm long,
conspicuous, coriaceous, taller than the corolla tube, elobate, opaque, epunctate,
glabrous, the apically free portions of the filaments terete, thicker than the

78 Sipa 18(1)

anthers, 0.8—-1.6 mm long, ventrally recurved at anthesis, epunctate, gla-
brous, the anthers subglobose, ca. 0.3 mm long, 0.4—0.5 mm wide, apically
rounded, basally cordulate, dorsifixed near base so as to appear basifixed,
the connective prominently red or black punctate dorsally; pistillode nor-
mally absent, occasionally conic, to 1 mm long, 0.3 mm wide, densely translucent-
lepidote. Pisti/late inflorescence resembling staminate in all features, but smaller,
2.5—3.5(-13) cm long, 3.5—8.0(—12.5) cm wide; peduncle 0.5—-1.5 cm long;
inflorescence branch bracts 3.0—3.5(—5.0) mm long, to 0.6 mm wide, at
times somewhat cucullate; floral bracts 2.0—2.6 mm long, ca. 0.5 mm wide,
pedicels virtually obsolete, or cylindrical to 0.1 mm long in flower, accrescent
and incrassate to 2.0 mm long, 1.0—1.5 mm diam. in fruit. Pisti//ate flowers
as in staminate, forming a condensed spike on the inflorescence branches;
calyx subcupuliform, 1.2—1.4 mm long; the tube to 0.1—0.2 mm long, the
lobes 0.9-1.1 mm long, 0.9-1.0 mm wide, widely ovate, apically acute;
corolla rotate, 2.7—2.9 mm long, the tube 0.6—0.7 mm long, the lobes ob-
long to elliptic, 2.0—2.2 mm long, 0.9-1.0 mm wide, apically acute; staminodes
1.5—1.6 mm long, the tube 1.0—-1.1 mm long, the apically free portions of
the filaments 0.4—0.5 mm long, the antherodes malformed, 0.2—0.3 mm
long, 0.3—0.4 mm wide; pistil obturbinate, 1.5—1.6 mm long, densely trans-
lucent-lepidote and prominently pellucid punctate, the style obsolete, the
stigma capitate, 0.2—-0.3 mm long, subsessile, 4-many-lobed, the lobes 1.3—
2.3 mm long, viscid, bright crimson, the placenta ovoid, the ovules 2, born
on side of placenta. Frit subglobose, 5.0O—9.0 mm long, 9-14 mm diam.,
fleshy, the exocarp thick, orange at maturity, prominently black punctate.
Bisexual Inflorenscence resembling staminate in all features except: 3.7—14.5
cm long, 2.5—-12.5 cm wide; inflorescence branch bracts and floral bracts
and pedicels as in pistillate. Bisexual flowers spicate, less crowded than in
the pistillate, more crowded than staminate; calyx cotyliform, (4—)5-parted,
1.0-1.2 mm long, the tube ca. 0.2 mm long, the lobes narrowly ovate to
narrowly triangular, 0.8-1.0 mm long, 0.3-0.4 mm wide, apically acute
to narrowly acute; corolla 4-lobed, rotate, 2.0—2.7 mm long, the tube ca.
0.3 mm long, the lobes oblong, 1.7—2.4 mm long, 0.9-1.2 mm wide, apically
acute; stamens 1.2—1.5 mm long, the tube ca. 0.5 mm long, the apically
free portions of the filaments 0.6-0.7 mm long, the anthers oblate, 0.3—
0.4 mm long, ca. 0.5 mm wide; pistil as in pistillate flowers except 1.2—-1.4
mm long, the ovary 1.0—1.1 mm long, 0.7—0.9 mm diam., the style short,
0.1—0.2 mm long, the stigma capitate, to 0.2 mm long, the lobes 0.9-1.1
mm long, the placenta ellipsoid, the ovules 2—3, borne on side of placenta.
Bisexual fruit suabglobose, as in pistillate, but 4-5 mm long, S—7 mm diam.

Distribution.—From Darién, Panama to Loreto, Peru, from 0—1,700 m
elevation.

Ecology and conservation status. —Cybianthus sprucei is a ridgetop species,

PrpoLy, Cybianthus in Ecuador and Peru 79

occurring in premontane wet and rainforests and also in lowland forests on
forest margins of exposed hilltops. This species, as opposed to its closest
congener, Cybianthus simplex, occurs in areas of high incident light for at
least part of the day. Because Cybianthus spruce continues to reproduce in
spite of mild forest intervention, it is not considered threatened or endan-
gered.

Etymology.—This species was named in honor of Richard Spruce, ardent
collector and student of the Andean and Amazonian flora.

Local names and uses.—Colombia: “margoandre,” “Tunda” Spanish (Valle
del Cauca, Colombia); Ecuador: “urcu tahucu” (Quichua); ground and a vapor
bath is taken to “send evil away” and to treat body pains, headaches, inter-
nal colds; Peru: “kurtp” Vivaro); the root is mashed and boiled, and the
decoction is drunk to “strengthen” the body.

Specimens an PANAMA. Darién: Cerro Pirré, 10— ye Loy (bisex fi tr).
Folsom 4544 (MO, PMA); on ridge of Cerro Pirré, 08° 00' N, 77° 45' W, 1,000-1,080 m,
14 Sep 1989 ae G. McPherson 14066 (BRIT, MO). EStoE Cauca: Rio Mice en
Guayabal, 5—20 m, 25 Feb 1943 (pist. fr), J. Cuatrecasas 14138 (COL, F, ae Dei Cauca,
El Tambo, 900 m, Apr 1937 (stam. fl), K. von Sneidern 1615 (S). Chocé: Along Rd. be-
tween Quibdé and Medellin, Km 207.5, 0-200 m, 18 Dec 1980 (pist. fl, fr), T; Croat G A.
re 52257 (COL, JAUM, MO); Alto del — 1,200-1,800 m, 8 Jan 1973 (pisc. fl, fr),

y G E. Forero 7317 (COL, F, MO). Narifio: La Guayacana, Funes, 24 Jun 1

(stam. aR R. Castaheda 2873 (COL, F); Ricaurte, 7 300 m; 18 Apr 1941 (bisex. fl), K. von
RSs A612 bis (S); Reserva Natural La Planada, 7 km above Chucunés on Rd. between
Tuquerres and Ricaurte, along Sendero La Vieja, 01° 06' N, 77° 54' W, 1,780-1850 m, 7
Mar 1990 (fr), T. Croat 71155 (MO, PSO). Valle del Cauca: Pacific coast, Rfo Naya, Puerto
Merizalde, 5—20 m, 22 Feb 1943 (bisex. fl), J. Cuatrecasas 14053 (COL, F); Rio Yurumangui,
550 m, 28 Jan—10 Feb 1944 (pisc. fl, fr), J. Cuatrecasas 15743 (COL, F, US); Rio Calima,
Chocé region, La Trojita, 5-50 m, 20 Feb 1944 (stam., pist. fl-sheets mixed), J. Cuatrecasas
16272 (COL, F, US); Rio Cajambre, Barco, 5-80 m, 30 Apr 1944 (fr), J. Cuatrecasas 17625
(COL, FE, US); Bajo Calima, 15 km N of Buenaventura, Cartén de Colombia concession,
aes be 03° 56'N, 77° 08' W, 500 m, 27 Mar 1986 (stam. fl), A. Gentry et al.
53713 (COL, MO), Concesién Pulpapel/Buenaventura, 03° 55' N, 77° 00' W, 100 m

Mar 1985 (fe), M. eae 767 (COL, CUVC, MO), 19 Mar 1985 (stam. fl), a Mansa
790 (CUVC, MO), 21 r 1985 (stam. fl), M. Monsalve 797 (CUVC, MO), 1

(stam. bud), M. ae ar (CUVC, MO), 24 Aug 1°86 (fr), M. Monsalve err cme
MO); Bahia de Malaga, near mouth of Quebrada La Sierpe,0 4° 00' N, 77° 15' W, 0-20 m,
17 Feb 1983 (stam. fl), A. Gentry et al. 40453 (COL, MO); 18 km E of Buenaventura, 50
m, 14 Feb 1939 (fr), E. Killip & H. Garcfa 33279 (BM, COL, NY, US), Buenaventura, Jun
nae an fs ne Lehmann B. T. 651 (K, NY). Putumayo: Umbria, 00° 54' N, 76° 10!

m, Jan—Feb 1931 (stam. fl), G. K/vg 2108 (US). ECUADOR. Bolivar : Monghest

a we ee El Tambo, 2,400 m, 18 Jul 1991 (stam. fl), H. van der Werff et al. 12430
(BRIT, MO, QCNE); along Rd. Chillanes-San Pablo, 6 oe hai Cai snes; 2,600 m, 21
Jul 1991 (fr), H. van der Werff et al. 12561 (MO, QCNE). Morona-Santiago: 15 km N of
Macas, Rd to Rio Upano, 02° 07'S, 78 °08' W, 1250 m, 5 Feb 1987 eee fl), J. Bohlin
et al. 1493 (GB); Cordillera de Cutucu, W slopes along trail from Logrono to Yaupi, 02°
46'S, 78° 06' W, 1,200 m, 10 Nov 1976 (stam. fl), M. Maddison et al. 3204 (US). Napo:
Canton Archidona, Carretera Hollin-Loreto, Rio Huataraco, 2 hrs walk from Guagua Sumaco,

80 Sipa 18(1)

00° 43'S, 77° 32' W, 800—-1,000 m, 23-30 Aug 1989 (stam. fl), C. Cerén & M. Factos
7648 (MO, QCNE); Cantén Orellana, Reserva Florfstica El Chuncho, 5 km N of Coca, 00°
25'S, 77° O1' W, 250 m, 23 May 1993 (fr), W. Palacios 10680 (MO, QCNE), El Chuncho,
el Payamino, Estacion sy elas INIAP-Napo, 5 km NW of Coca, 00° 30'S, 77° O1
W, 250 m, 12 Oct 1987 (stam. fl), C. Cerén 2494 (MO, QCNE); Cantén Tena, Mishualli,
junction of Rios Mishualli oa Napo,01° 03'S, 77° 41' W, 500 m, 13-14 Aug 1979 (fr),
L. Holm-Nielson 19295 (AAU), Estaci6n Biolégica Jatun Sacha, along S bank of Rio Napo,
1 km E of Puerto Misahualli, 00° 04'S, 77° 36' W, 450 m, 1 Apr 1992 (fr), T) Croat 73352
(MO, QCNE), 8 km E of Puerto Mishualli, 01° 04'S, 77° 36' W, 400 m, 14 Aug 1989
(ster.), C. Cerén 7409 (MO, QCNE), 22 Sep 1989 (stam. fl), W. Palacios 447 1 (MO, QCNE),
16 Mar 1991 (fr), D. Nez// 9813 (BRIT, F, MO, QCNE); Rio Blanco Comunidad, penn
ters of Rio Huambuno, 6 km NNW of Ahuano, 01° 44' §S, 7 44' W, 440 m, 18 Jul—
Aug 1990 (tr), E. Kohn 1311 (MO). PERU. Amazonas: Prov. Bagua, Yamayakat, me
Putuim, 04° 55'S, 78° 19' W, 500 m, 17 Oct 1996 (stam. fl, R. Vasquez & N. Jaramillo
20318 (AMAZ, BRIT, MO). Loreto: Prov. Loreto, Pampa Hermosa and vicinity, Rio Corrientes,
| km S of junction with Rio Macusari, 03° 15'S, 75° 50' W, 160 m, 3-20 Dec 1985 (stam.
fl), W. Lewis et al. 10180 (BRIT, MO, USM).

Cybianthus sprucei was misinterpreted by Mez, and confused with C. szm-
plex (Mez 1902). From there, Werge/tia panamensis was described based pri-
marily on subtle differences and geography. Finally, Cuatrecasas described
Weigeltia purpurea from the Chocé floristic region of Colombia, notable only
for its narrower leaves, the secondary veins more arcuate, and some quanti-
tative floral characteristics

Cybianthus sprucei is most closely related to C. simplex because of the homomerous
flowers, long sa es and non-mucronulate leaf apices. However, Cybianthus
sprucei may easily be separated from C. simplex because of its arborescent
habit, pyramidal panicle with spicate branches, symmetric leaf base and longer
cataphylls.

19. aera simplex (Hook. f.) G. Agostini (Fig. 4C), Acta ere
10:163. 1980. Comomyrsine simplex Hook. f. in Benth. & Hook., Gen 2:644,
1876. Pe ane Mt AG f.) Mez ‘v Engl., Pflanzenr IV. 236(Heft 9): on 1902.
Type: ECUADOR. Cuimporazo: At foot of Volcin Chimborazo, 760 m, Aug 1860
ae fl, fr), R. Spruce 6143 (HOLOTYPE: K-2 sheets).

—_—

oo Diels, Notizbl. Bor. Gart. Berlin-Dahlem 15:383. 1941. ee nov.

nthus sels (Diels) G. see Acta Biol. Venez. 10:163. 1980 PE:
a : Mera, 1,200 m, 15 Nov 1938 (stam. fl), H. Schultze- ca
2983 (HOLOTYPE: B des estr.; no isotype found). I defer neotypification until material

has been regathered from the type locality or an adjacent one

Semi-woody subshrub to \(—3.5) m tall. Stem terete, 1-5 cm diam., sparsely
glandular-papillate, glabrescent. Cataphyl/s few, spirally arranged in inter-
nodal areas, coriaceous, subulate, 6.5—2 m long, 1.12—1.7 mm wide,
keeled, densely and prominently Blade aes and punctate-lineate, sparsely
glandular-papillate, the margin flat, entire. Leaves pseudoverticillate; blades
membranaceous, elliptic to oblong, (34.5—)38-46.5(—80) cm long, (12.5-)

Pipoty, Cybianthus in Ecuador and Peru 81

13.5-18.5(—32.5) cm wide, apically broadly acute or rounded to a short
acumen, rarely acute, basally asymmetric,broadly rounded or rarely abruptly
acute, slightly decurrent on the petiole, hydropotes few above, numerous
below, often sparsely glandular-papillate below, conspicuously black punc-
tate, the margin irregular, hyaline when juvenile, opaque at maturity, flat,
entire; petioles rigid, deeply canaliculate, 7—17(—21) cm long, 4—6 mm diam.,
slightly marginate at junctions of the blade, expanded basally and slightly
decurrent on stem, sparsely glandular-papillate, glabrescent. Inflorescence bracts
membranaceous, linear-lanceolate, 17-25 mm long, 2-4 mm re apically
long-attenuate, hyaline, densely and prominently black punctate-lineate,
the margin irregular, entire; peduncle (1.2—)2.44.5 cm long. Staminate inflorescence:
a thyrsoid panicle (columnar) 11—28(—54) cm long, bi- or tripinnate, the
primary branches subopposite, each branch pyramidal-paniculate, the flowers
racemose; inflorescence branch bracts linear-lanceolate, subulate, (SO—)61—
72 mm long, 0.5—-1.3 mm wide, apically long-attenuate, medially keeled,
hyaline, glandular-papillose throughout, the margin entire; floral bracts early
caducous; pedicels 2.3-3(—5) mm long, glandular-papillate and -ciliolate.
Staminate flowers 5-merous, green to greenish-white when fresh; calyx mem-
branaceous, subcotyliform, 1.1—1.4 mm long, equally divided, the tube 0.2
mm long, the lobes lanceolate to lanceolate-subulate, apically narowly acute
to long-attenuate, hyaline, epunctate, the margin erose-dentate; corolla coriaceous,
rotate, 2.2—3.1 mm long, the tube 0.3—0.7 mm long, the lobes linear-lan-
ceolate, 1.5—2.6 mm long, 0.9-1.1 mm wide, reflexed distally 180° from
the tube, the apically subulate, densely glandular-granulose throughout within
and along the margin, epunctate, the margin regular, entire; stamens |.0—
2.3 mm long, the tube conspicuous, coriaceous, 0.3—0.7 mm long, the apically
free portions of the filaments terete, as wide as the anthers, 1.0-1.1 mm
long, ventrally recurved, the anthers connivent at first, subglobose, OZ
0.5 mm long, 0.5—0.6 mm wide, apically rounded, basally cordulate, dorsiftxed
just above the base, the connective prominently black punctate dorsally;
pistillode absent. Pzstillate inflorescence: as in staminate but 16—30 cm long;
inflorescence branch bracts ovate-subulate, 4.5—7 mm long, 1.5—2 mm wide,
the margin irregular, erose; floral bracts membranaceous, linear-subulate,
1.6—2 mm long, 0.4—0.6 mm wide, apically long-attenuate, densely glan-
dular-papillose puberulent, the margin irregular-entire; pedicel terete, 1—
10 mm long, translucent, glandular-puberulent. Pisti//ate flowers green, ca-
lyx coriaceous, 0.6—1 mm long, unequally divided, the tube 0.2—0.3 mm
long, the lobes ovate to widely ovate, 0.4—0.7 mm long and wide, apically
acute, often moderately glandular-papillate without, the margin irregular,
erose; corolla 1.5—1.7 mm long, the tube 0.4—0.5 mm long, the lobes ob-
long, 1.0—-1.2 mm long, 0.7—0.8 mm wide, apically acute, reflexed 135°

82 Sipa 18(1)

from tube, glandular-papillate without; staminodes ().8—1.2 mm long, the
tube inconspicuous, membranaceous, 0.4—0.5 mm long, glabrous, elobate,
the apically free portions of the filaments 0.3—0.4 mm long, the antherodes
subglobose, 0.2 mm long, 0.3 mm wide, apically rounded, basally cordulate;
pistil subglobose, 0.8—1.1 mm long, the ovary 0.6-0.8 mm long, 1.0-1.2
mm diam., densely translucent-lepidote, the style short, thick, 0.1 mm long,
0.7 mm diam., the stigma capitate, the margin lacinate, with numerous
lobules to 0.2 mm long, the placenta umbonate, the ovules 3, exposed apically
1/3 their length on the placenta. Fruit subglobose, 4-5 mm long, 5-8 mm
diam., red, exocarp somewhat thick, juicy, prominently black punctate.

in Colombia and Ecuador, 60—2,200 m.

Ecology and conservation status. —Cybianthus simplex occurs in premontane
wet and rainforests, occasionally in the understory of ridgetops forests. This
species occurs in deep shade under the shrub stratum of the forest. It occurs
more frequently i in primary forest wet enough to maintain Chusquea popu-
lations, but can survive in disturbed forests as long as the shrub and Chusquea
populations exist. Because of its habitat flexibility, it is not considered threatened
or endangered.

Etymology.—The epithet “simplex” refers to the monoaxial habit of the
species.

Specimens examined. COLOMBIA. Narifio: Mpio. Barbacoas, Corregimiento Ortiz y
Zamora, Vereda E] Barro, Reserva Natural Rio ee ca. 5 km W de Altaquer, faldas
occidentales de la Cordillera Occidental, 01° 18' N, 2 08! W, 1,350-1,400 m, 1 Sep

1997 eee fl), J. Pipoly, A. Cogollo, et al. 21026 , ee 21051, 21055 (BRIT, FMB,
JAUM, PSO), 2 Sep 1997 (bisex. fl), J. Pipoly, A. —o et al. 21109(BRIT, JAUM, PSO),
(ster.) J. Pzpoly, A. Ue 21131, 21148 (BRIT, PSO), ADE fl, fr), J. Pipoly, A. Cogollo
et al. 21173 (BRIT, FMB, JAUM, PSO), 1,490- : a m, 4 Sep 1997 (stam. fl), J. Pépoly,
A, Cogollo et al. ee (BRIT, FMB, JAUM, PSO), (pist. fl), J. Pzpoly, A. a et al. 21296
(BRIT, FMB, JAUM, PSO), 1,350—1,400 m, 5 Sep 1997 (stam. fl), J. Pipoly, A. Cogollo et
al. 21326 (BRIT, JAUM, PSO), (ster.), J. Pipoly, A. Cogollo et al. 213 - ie JAUM,
PSO),1,350-1,145 m, 6 Sep 1997 (ster.), J. Pipoly, A. Cogollo et al. 21417 (BRIT, FMB,
JAUM), 7 Sep 1997 (tr), J. Pipoly, A. Cogollo et al. 21469, 21471, 1,450-1,500 m, 8 Sep
1997 (stam. fl), J. Pipoly, A. Cogollo et al. 21520 (BRIT, FMB, JAUM, PSO), (ster.), J. Pipoly,
A. Cogollo et al. 21524 (BRIT, JAUM, PSO), along trail from main Pasto-Tumaco Rd. to
Rio Nambj, departing main Rd. at ba Mixta El Mirador, 7 km W of Altaquer, 01°
18'N, 78° 04' W, 1,100 m, 26 Feb 1992 (fr), T. Croat 72394 (JAUM, MO); Corregimiento
ger Vereda el Barro, Reserva Natural Rfo Nambi, W slope, W Cordillera, 01° 18'

° 08' W, near Cabafia Fundacién FELCA, 1,325 m, 11 Dec 1993 (fr), J. Betancur et al.

ae (COL, MO); La Planada, Finca Salazar, 7 km above Ricaurte, 01° 08' N, 77° 58' W,

1,750 m, 29 Nov 1981 (pist. fl, fr), A. Gentry et al. 35188 (BRIT, COL, MO, US); La Planada,

S ee ars 7 km from Tumaco-Pasto Rd., 01° 10'N, 77° 58' W, 1,800 m, 24 Jul 1986

. m. fl), A. Gentry et al, 55053 (MO, PSO); trail to Hondén, 6-12 km SW of La Planada,

04' N, 78° 02' W, 1,750-1,800 m, 5 Jan 1988 (fl bud), 0. de Benavides & R. ae
yee (MO, PSO); Valley of Rio Guiza, Rd. from El Espino to Tumaco, ca. 21 km

Prpoty, Cybianthus in Ecuador and Peru 83

Ricaurte, 01° 15'N, 78° 07' W, 1,000 m, 7 Dec 1988 (stam. fl), B. Hammel 17150 (JAUM,
MO). ECUADOR. Bolivar: Hacienda Changuil, LA 16; 02° 06'S, 79° 10' W, 500 m, 17
Aug 1995 (stam. fl), X. hae Bonifaz 4339 (GUAY, MO). Gihe Prominent hillcrest
directly N of Lita, on N side of Rio Mira, E of Rio Baboso, W-facing slope, 00° 53' N, 78°
27' W, 760 m, 7 Aug 1994 eon B. Boyle 3473 (MO, QCNE), Steep N-facing slope S of
Baboso, S side of Rio Baboso, 00° 53' N, 78° 27' W, 750 m, 11 Aug 1994 (ster), B. Boyle
3599 (MO, QCNE); Rio Blanco drainage above Chical, tributary of Rio San Juan, 12 km
W of Maldonado, 1300-1500 m, 25 Sep 1979 (bisex. fl), A. Gentry & G. Shupp 26565
(MO, QCNE); Cantén Tulcén, Parroquia Tobar Donoso, Reserva Indfgena Awa, Centro El
Baboso, 00° 53' N, 78° 25' W, 1,800 m, 17-27 Aug 1992 (stam. fl), G. Tipaz et al. 1709
(BRIT, MO, QCNE), (stam. fl), G. Tipaz et al. 1886 (MO, QCNE), (fr), G. Tipaz et al. 1924
(MO, QCNE); 6 km above Maldonado, just below Puente de Palo, 00° 54' N, 78° 06' W,
2,275 m, 23 May 1993 (stam. fl), B. Boyle GJ. Bradford 1878 (MO, QCNE); cae ae
Paflon to Gualpi Chicé, Reserva Indfgena Awa, 1.5 km past Rio Blanco, 00° 51'N,
16' W, 1,000-1,450 m, 14 Jan 1988 (stam. fl), W. Hoover et al. 2456 (MO, ene
Trail, Gualpi Chic6 area of Awa Reserve, 00° 58' N, 78° 16' W, 1,330 m, 19 Jan ce
(pist. fl, fr), W. Hoover et al, 2809 (MO, QCNE); Trail along ridge and forest slope to NW
of oe encampment, Gualpi Chic6 area near Finca Rodriguez, 00° 58' N, 78° 16' W, 1,258-
1,323 m, 19 Jan 1988 (fr), W Hoover et al. 3358 (MO, QCNE). Coropaxt: Rio Guarapa, ca.
20 en NW of El Corazén, 250 m, 19 uit 1967 (stam. fl), B. Sparre 17091 (S), 20 Jun
1967 (pisc. fl), B. Sparre 17081 (S). El Oro: 11 km W of Las Pifias on new Rd. to Sta. Rosa,
850 m, 8 Oct 1 ee ee fl), C. Dodson et al. 9101 (MO, SEL); Hacienda Buenaventura,
12 km W of Las Pifias on Rd. to Machala, 03° 48'S, 79° 46' W, 1,000 m, 1 Mar 1991
(stam. fl), M. ee ve (GOET, MO); New Rd. Saracay-Balzas-Velacruz, ca. 8 km SE of
Saracay, 400 m, 30 Apr 1980 (stam. fl), G. Harling & L. Andersson 18778 (GB). Guayas:
Cordillera Chog6n-Colonche, Cerro Los Pontones; 01° 44'S, 08° 40' W, 500 m, 2 Jul 1994
(stam. fl), X. Cornejo & C. Bonifaz 2979 (GUAY, MO). Loja: Tierra Colorada, 1 km E of
Landara, 8 km E of Mercadillo, 04° 02'S, 79° 57' W, 1,500 m, 9 Feb 1991 (pist. fl, fr), M.
Kessler 2401 (BRIT, GOET). Manabi: Machalilla National Park, zona de San Sebastian,
01° 36'S, 80° 42' W, 600-700 m, 21 Jan 1991 (fr), A. Gentry et al. 72499 (MO, QCNE)
Pichincha: Quito-San Juan Chibore: Sto. Domingo de los Colorados Rd., Branch km
59, 18 km NW of Rd., 1,700—2,000 m, 27 Sep 1986 (bud), V. Zak 1350 (MO, US); Quito-
Aloag-Sto. Dominigo de los Colorados, km 94, 10 km S of Rd., W slopes of Volcan El
i 00° 21' 30 “S, 78° 51' 15” W, 1,300-1,500 m, 25 Dec 1986 (fr), V. Zak 1545
(MO, US); 15 ha. Patch of forest in Cooperativa Sta. Marta No. 2, along Rfo Verde, 2 km
SE of Sto. Domingo de Los Colorados, 530 m, 5 Feb 1979 (fr), C. Dodson et al. 7597 (MO,
SEL); Reserva Floristica-Ecolégica “Rio Guajalito,” Km 59, Quito-Santo Domingo de los
ea 3.5 km NE of Rd., lower slopes of Volcan Pichincha, 00° 13'53"S, 78° 48' 10"
00-2,200 m, 28 Dec 1985 (fr), J. Jaramillo 8298 (MO, QCA), Cantén Quito, pees

>

a Reserva eae anne ee Inca Trail to Rio Tulambj, ca. 5 airline km
ee 00° 07' N, 78° 38' W, ease m, 15 Sep 1989 (fr), G. Webster & P. Delprete aay
(DAV. A), along trail between Rfo Umachaca and Rio Tulambi, 00° 07.5' N, 78° 38.5!

» QC

W, 1,200-1,300 m, 7 Jul 1990 (fr), 2 Webster et al. 27795 (DAV, QCA); Montafias de
pee see Cerro Sosa, 00° 0.5' N, 78° 37' W, 1,950 m, 3 Jul 1991 (fr), G. Webster
28702 (DAV, QCA), 1,750 m, ae ae (stam. fl), G. Webster et al. 28710 (DAV, A),
on ridge heeween Base Camps | & 2, 00° 5.5' N, 78° 37' W, 1,800-1,900 m, e 7 Jul

1991 (fr), G. Webster & B. Castro 28769 (DAV, MO, QCA); along Rio Umachaca near Ha-
cienda El Carmen, 00° 07-7.5' N, 78° 38' W, 1,250 m, 6—7 Jul 1991 (fr), G. ee
28796 (DAV, QCA). Quininde: Bilsa Biological Reserve, Montafias de Mache, 35 km
of Quinindé, 5 km W of Sta. Isabela, SE ridge trail, 00° 21' N, 79° 44' W, 400-600 m,

84 Sipa 18(1)

Sep 1994 (stam. fl), N. Pitman et al. 688 (MO, QCNE), : <i aye (fr), N. oF 993

(MO, QCNE), Along Dogala and a Trails, OO° 2 'W/, 400-600 m, 2

: in 1995 (fr), N. Pitman 1161 (M CNE). Province ee n. are os
1896 (pist. fl), J. Sediro 100/12 is a F Neg 4859).

This species has often been confused with Cybianthus spruce, owing to
variation in leaf morphology and inflorescence size. However, recent field
studies conducted at the Rio Nambi Natural Reserve of Narifio, Colombia,
have revealed that juvenile individuals have obtuse to somewhat broadly
rounded, asymmetric leaf bases with long petioles, while mature individu-
als have tapering, asymmetric leaf bases. The confusion was due, in part, to
precociously flowering individuals, detectable by their extremely small flowers,
or to reiterative shoots, detectable by the renewal shoot visible below the
“bayonet”, that bears juvenile leaves and pink flowers. While the largest
individuals of Cybianthus simplex may approach the size of many C. spracei,
the large pith of the stem in the former renders them extremely weak, and
the stems may easily be snapped by hand, while the pith of C. spracez is
relatively smaller, and the stems can be bent without snapping in the field.

Cybianthus simplex is most closely related to C. sprucer, but may be sepa-
rated from it by the columnar, thyrsoid panicles with racemose branches,
the pedicellate flowers, asymmetric leaf base and shorter cataphylls. Popu-
lations corresponding to the type of Weigeltia chamaephyta differ from the
type of C. smplex in floral structure, directly attributable to the fact that

the former is based ona pistillate, and the latter a staminate collection. However,

the autapomorphic columnar thyrsoid panicle leaves no doubt that they are
synonymous. Because no further collections have been made in the region
from which the type was collected, I defer neotypification until collections
from that area are available.

20; oo kayapu (Lundell) Pipoly, comb. nov. (Figs. 3E, 4F, 6J,
A-F). Weigeltia kayapii Lundell, Wrightia 6:118. 1980. Type: PERU. AMAzonas:
mino de chichijam, entsa, 300 m, 2 May 1973 (fr), R. Kayap 723 (uoLoryPe: MO;

isoTYPE: LL-TEX).

Monoaxtal treelet to \(—2) m tall. Stems terete, (0.6—)1—1.7 cm diam., glandular-
papillate at first, glabrescent. Cataphy//s alternate in a high spiral, coria-
ceous, subulate, 15—45 mm long, 0.5—2(—3.5) mm wide. Leaves pseudoverticillate,
blades chartaceous, oblanceolate to oblanceolate-oblong, (22—)31.3—55.5
cm long, 8.5-19.6(—23) cm wide, apically acute or broadly rounded to a
small acutish tip, mucronate, the mucron to 0.5 mm long, the blade gradually
tapering to an abruptly obtuse base appearing auriculate, to 1.5 cm wide,
midrib slightly raised above, prominently raised below, the secondary veins
12-16 pairs, with prominent marginal and submarginal collecting veins,
slightly sunken above, prominently raised below, glabrous above, with ru-
fous hydropoten below; petiole deeply canaliculate, 1-2 cm long, ca. 3.5—

PipoLy, Cybianthus in Ecuador and Peru 85

4 mm diam., densely glandular-papillate adaxially. Staminate inflorescence: a
pyramidal, pipianae panicle, (3—)5.5—29 cm long, 5—15(—26) cm wide, the
branches racemose, densely glandular-papillate, succulent, then drying hyaline;
peduncle 3—5.5 cm long; branch bracts membranaceous, subulate, 6.5—8
mm long, 0.5—1.5 mm wide; pedicels cylindrical, 1.2—1.8 mm long, sparsely
glandular-papillate, glabrescent; floral bracts membranaceous, subulate, inserted
on the pedicel about at middle, longer than the pedicel, 1.5—2.5 mm long,
0.1—0.2 mm wide, hyaline, densely glandular-papillate, the margin entire.
Staminate flowers pink, heteromerous, the calyx 5-merous, the corolla 4-merous;
calyx deeply membranaceous, cupuliform, 0.9—1.1 mm long, the tube 0.1—
0.2 mm long, unequally divided, the lobes deltate to subdeltate, 0.6—-0.9
mm long, 0.2—0.7 mm wide, highly reflexed at anthesis, apically acute, epunctate,
hyaline, densely glandular-papillate, the margin glabrous, entire; corolla
carnose, subrotate to rotate, 2-3 mm long, the tube 0.5—0.8 mm long, the
lobes ovate, 1.5—2 mm long, 1.1—1.6 mm wide, apically acute, distally re-
curved 90° from tube axis at anthesis, opaque, densely glandular-granulose
within and along margin, sparsely glandular-papillate along margins without,
epunctate or sparingly and inconspicuously pellucid punctate, the margin
entire; stamens 2.2—2.9 mm long, subequal to corolla lobe or exserted, the
tube conspicuous, carnose, 0.5—-0.8 mm long, hyaline, glabrous, elobate,
the apically free portions of the filaments terete, 1.6—2.2 mm long, free from
corolla, proximally recurved, the anther oblate, 0.3-0.5 mm long, 0.5—0.8
mm wide, always wider than long, apically emarginate to retuse, basally
widely cordate, the connective prominently black punctate dorsally, con-
spicuously black punctate ventrally; pistillode absent or to 1 mm long, 0.1—
0.3 mm wide, densely glandular-papillate. Pisti//ate inflorescence as in stam1-
nate but 6.5—9(—10.5) cm long, erect, not succulent, opaque, densely
glandular-papillate; peduncle 1-2 cm long; branch bracts 2-3 mm long,
0.20.3 mm wide; pedicels subobsolete or cylindrical, to 1.2 mm long, incrassate
and accrescent in fruit to 1.5 mm long; floral bracts inserted on pedicel,
longer than the pedicel, 1—1.3 mm long, 0.2—0.3 mm wide. Pzstillate flowers
as in staminate but pink to pinkish-white; calyx 0.9-1.2 mm long, the lobes
unequally divided, the smaller linear-lanceolate, 0.8—0.9 mm long, 0.3—
0.4 mm wide, the larger deltate, 1.0—1.1 mm long and wide; corolla ro-
tate, 2.6—2.9 mm long, the tube 0.9—1 mm long, the lobes elliptic, 1.7—
2.0 mm long, 0.6—0.7 mm wide, reflexed at anthesis, distally recurved 180°
from tube axis, glabrous without, sparsely glandular-granulose within, the
margin slightly irregular; staminodes very poorly developed, 1.2-1.5 mm
long, the tube 0.9—1.0 mm long, the apically free portions of the filaments
(.3—0.6 mm long when developed, recurved proximally, the anthers badly
formed, at times consisting of 2—3 prominently punctate glands at filament
apex, or otherwise as in the stamens, ovate to subglobose, 0.2—0.3 mm long,

86 Sipa 18(1)

0.3—0.4 mm wide, apically irregular, obtuse, emarginate or retuse, basally
cordate, the connective when distinguishable prominently black punctate
dorsally, conspicuously punctate ventrally; pistil clavate to lageniform, 3—
3.5 mm long, the ovary 0.9-1.1 mm long, 1.2—1.5 mm diam., densely papillate,
the style 2.1—2.4 mm long, the stigma large, capitate, with 4 principal lobes,
each irregularly lacinate-lobulate, early caducous, the placenta deeply cupuliform,
the ovules 2, buried for 1/2 their length. Frit depressed-globose, 5—6 mm
long, 7-9 mm wide, prominently black punctate, the exocarp thin. Bisexal
inflorescence: as in staminate but 4-13 cm long. Bisexual flowers as in stami-
nate flowers but calyx 1.1—1.9 mm long, the tube 0.2—0.3 mm long, the
lobes unequally divided, deltate to elliptic, the smaller 0.6-0.7 mm long
and wide, the larger 0.9-1.6 mm long, 0.5—0.6 mm wide, otherwise as in
pistillate flowers; corolla 2.6-2.8 mm long, the tube ca. 0.6 mm long, the
lobes narrowly ovate, 2.0—2.2 mm long, 1.2—1.3 mm wide, recurved dis-
tally 90° from tube, sparsely glandular-granulose within, glandular-papil-
late along the margin; stamens as in staminate flower, but 2.2—2.7 mm long,
always slightly shorter than corolla tube, the tube ca. 0.6 mm long, the
apically free portions of the filaments 2.0—2.2 mm long, the anthers widely
ovate, ca. 0.3 mm long, 0.5—0.6 mm wide, apically obtuse to emarginate,
basally widely cordate; pistil 2.7-2.9 mm long, the ovary 0.8—0.9 mm long,
0.6—0.7 mm diam., densely glandular-papillate, the placenta deeply cupuliform,
the ovules 2, buried for 1/2 length. Bisexual fruit depressed-globose, 5—6
mm long, 6—7 mm wide, prominently black punctate, the exocarp thin.

Distribution.—Colombia (Amazonas, Chocé, Narifio, Valle del Cauca, ),
Ecuador (Chimborazo, Napo and Pichincha), Peru (Amazonas, Loreto, San
Martin) and Brazil (Acre), growing at sea level—2,530 m elevation.

Ecology and conservation status. —Cybianthus kayapii is locally common in
small populations at the high water line in primary “tahuampa” habitats
(varzea forest), along white water rivers, or rarely in premontane habitats
along the edge of pools fed by creeks. The species is not known to be culti-
vated and occurs only in areas where deep leaf litter and alluvial deposits
are left after flooding action. It appears that the species can easily be de-
stroyed by soil compaction as a result of trampling, and thus, it should be
considered threatened.

Etymology.—The epithet commemorates Rubio Kayap, an indigenous
Aguaruna Peruvian plant collector who worked with Brent Berlin, known
for his great knowledge of Amazonian flora and ethnobotany.

Local names and uses. —Colombia: “Hierba de palo grande”(Spanish). Ground
in crude syrups to cure cuts, internal infections, to clean the uterus and the
stomach. Ecuador: “putush” (Shuar). Used against intestinal parasites and
for chronic rectal bleeding (colo-rectal carcinoma °); 10 Ibs. of root boiled
in 8 liters of water, down to one liter; 8 cc given as enema before bed; useful

Prpoty, Cybianthus in Ecuador and Peru 87
for “dysentery.” Peru: “mantaya,” “kugkuima muspari” (Aguaruna); used
to disinfect dog and insect bites; “napi tsuake” (Huambisa). Brazil: used in

curare cf. B. Krukoff 7603.

Representative specimens examined. COLOMBIA. Amazonas: Mpio. Leticia, Parque
Nacional Natural Amacayacu, Centro Administrativo Mata-mat4, trail to Amacayacu, km
4, 03° 47'S, 70° 15' W, 120 m, 25 Sep 1991 (stam. fl), A. Rudas G A. Prieto 3147 (COL,
FMB, MO), 110-120 m, 28 Oct 1991 (fl bud), J. Pipoly & J. Murillo 15483 (COL, FMB,
MO), Quebrada de Agua Pudre, ca. 1.5 km NE of junction with Rio Amacayacu, perma-
nent inventory plot, 200-220 m, 11 Nov 1991 (ster.), J. Pipoly et al. 15896 (COL, FMB,
MO), 15 Nov 1991 (stam. fl), J. Pzpoly et al. 16075 (COL, FMB, MQ); Rio Loretoyacu, 100
m, Oct 1946 (bisex. fl), R. E. Schultes & G. Black 8427 (US). Chocé: Bahia de Solano, 13—
18 Apr 1982 (pist. fl, fr), R. Dressler 6036 (COL, FLAS, MO); Mpio. de Quibdé, Corregimiento
San Francisco Iché, Quebrada Caledonia along Caledonia Rd., 9 Apr 1987 (ster.), EF Garcia
& J. Echavarria 259-A (COL, CHOCO, MO). Narifio: Mpio. Ricaurte, Reserva Natural La
Planada, 1,800 m, 13 Nov 1993 (fr), C. Restrepo 723 (BRIT, MO, PSO); La Planada, Finca
Salaazar, 7 km above Ricaurte, 01° 08' N, 77° 58' W, 1,750 m, 27 Nov 1981 (fr), A. Gen-
try et al. 35062 (MO, PSO). Narifio: Mpio. Barbacoas, Corregimiento Ortiz y Zamora,
Vereda El Barro, Reserva Natural Rfo Nambjf, ca. 5 km W de Altaquer, faldas occidentales
de la Cordillera Occidental, 01° 18' N, 78° 08' W, 1,350-1,400 m, 3 Sep 1997 (ster.), J.
Pipoly, A. Cogollo, et al. 21241 (BRIT, FMB, JAUM, PSO). Valle del Cauca: Rio Calima,
region del Chocé, La Trojita, 5-50 m, 20 Feb 1944 (stam. fl), J. Cuatrecasas 16272 (COL,
F, US); Mpio. El Cairo, Cerro del Inglés, summit, Cordillera Occidental, Serrania de los
Paraguas, limit Valle/Chocé6, El Cairo-Rio Blanco Hwy, | hour in jeep from El Cairo, 2,400
m, | Jan 1987 (pist. fl), P. Silverstone-Sopkin et al. 2871 (CUVC). ECUADOR. Carchi: Cantén
Tulcaén, Parroquia Tobar Donoso, Reserva Indfgena Awa, Centro El Baboso, 00° 53' N,
78° 25' W, 1,800 m, 17-27 Aug 1992 (stam. fl), G. Tipaz et al. 1706 (MO, QCNE). Esmeraldas:
Cantén San Lorenzo, Parroquia Ricaurte, Reserva Indigena Awd, Comunidad Balsarefio,
Rio Palab/, 01° 09' N, 78° 31' W, 100 m, 15-29 Apr 1991 (fl bud), D. Rubio & C. Quelal
1335 (MO, QCNE). Chimborazo: Cordillera Occidental, “El Carmen,” Sibambe, 2,450
m, 22 Aug 1943 (pist. fl), M. Acosta-Solés 5544 (F, QCNA); on slopes of Chimborazo Vol-
cano, (pist. fl, fr), A. Sodrro 100/14 (B, destr., QA?, n.v.). Morona-Santiago: Basin of Rio
Morona, Rio Mangosiza, Nayumbime, 45 km SE of Sucua (by air), 200 m S of Don Luis
Najamte’s house, ca. 02° 43'S, 77° 38' W, 300 m, 27 Feb 1990 (fr), C. Limbach 140 (QCA,
NY). Napo: Antisana, Shinguipino Forest, between Rios Napo and Tena, 8 km SE of Tena,
01° 00'S, 77° 50' W, 450 m, 17 Sep 1960 (stam. fl), P. Grubb et al. 1633 (K, NY); Cantén
Archidona, Carretera Hollin-Loreto, Rfo Huataraco, 2 hrs by foot from Guagua Sumaco,
00° 43'S, 77° 32' W, 800—1,000 m, 23-30 Aug 1989 (fl bud), C. Cerin & M. Factos 7641
(MO, QCNE); Cantén Tena, 17 Oct 1939 (stam. fl), E. Asplund 9396 (S); 3 km E of Caserfo
Huamani, N of Carretera Hollin-Loreto, 00° 43'S, 77° 36' W, 1,200 m, 17 Sep 1988 (fr),
FE. Hurtado & A. Alvarado 503 (MO, QCNE). Pichincha: Along Rd. from Tandayapa to
Mindé, 10 km from Tandayapa, 2,530 m, 16 Dec 1979 (fr), T. Croat 49361 (MO, QCNA).
PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Comunidad Aguaruna de Kampaentza (PUJAIM),
property of Juan Maydn, 740 m, 6 Oct 1994 (pisc. fl), C. Diaz et al. 7265 (BRIT, HUT,
MO, USM); Prov. Condorcanqui, Drto. El Cenepa, NE region of Marafion Drainage Basin
Rio Cenepa, Comunidad Tutino, 04° 33'S, 78° 10! W, 350 m, 21 Nov 1993 (pist. fl, fr),
R. Vasquez et al. 18395 (AMAZ, BRIT, HUT, MO, USM); Rio aie vicinity Saeueyen
ca. 5 km E of Chavez Valdivia, 04° 30'S, 78° 30' W, Quebrada Chigkishinuk, 10 Apr
1973 (fr), E. Ancuash 211 (MO), 11 Aug 1978 (pist. fl), E. ae 1405 (MO); Trail one
day walk from Huampami to Shaim, creek running into Nahim, 600 m, 27 Nov 1972

88 Sipa 18(1)

Pic. 17. Cybianthus anthuriophyllus Pipoly. A. Habic, showing pleiochasium with sympo-
dial branches. B. Staminate flower, showing lanceolate-subulate corolla lobes, elobate staminal
tube, and conic, vestigial pistillode. C. Pistillate flower, showing attenuate-acicular calyx
lobes and obturbinate pistil. A, C, drawn from holotype. B, drawn from Bravo & Gomez 49.
Figure drawn by Peggy Duke.

Pipoty, Cybianthus in Ecuador and Peru 89

(stam. fl), B. Berlin 393 (MO); Rio Santiago, W bank, 400 m beyond La Poza, 180 m,
without date (fr), F Dominguez 147 (MO); 800 m beyond Caterpiza, 200 m, 4 Sep 1979
(fr), V. Huashikat 356 (MO), 10 Sep 1979 (stam. fl), Vi Huashikat 507 (MO), 12 Sep 1979
(bisex. fl), V. Huashtkat 581 (MO); 26 Mar 1980 (fr), 8. Tungué 1110 (MO). Loreto: Brey
Maynas, Explorama Lodge, near Yanamono, between. Indiana and mouth of Rio Napo 03
28'S, 72° 50' W, 103 m, 27 Jun 1983 io. Gentry et a 42247 (AMAZ, MO, NY), 106 m,
15 Apr 1985 (fr), R. Vasquez GN. oo oe 25 (AMAZ, MO), 28 Sep 1988 (stam. fl), R.
Vasquez & N. Jaramillo 11100 (AMAZ, M 1 Jul 1990 (fr), R. Vasquez & N. Jaramillo
14092 (AMAZ, MO, US), 25 km NE of aie along Rio ee 90 m, 03° 30'S,
72° 50! W, 26 Sep 1990 (stam. fl), J. Pipoly et al. 12383 (AMAZ, MO), along S border
trail, 110 m, 27 Sep 1990 (stam. fl), J. Pipoly et al. 12490 one MO, NY, US, USM),
(stam. fl), J. Pipoly et al. 12541 (AMAZ, MO, US, USM); Prov. Alto Amazonas, N slopes of
Cerros Camapaquiz at Pongo de Manseriche, right bank of Rio Marafion, 300-550 m, 19—
21 Oct 1962 (stam. fl), J. Wurdack 2324 (US, USM). San Martin: Pongo de Cainarachi,
Rio Cainarachi, tributary of Rfo Huallaga, 230 m, Sep-Oct 1962 (stam. fl-except speci-
men at S-bisex. fl), G. Klug 2691 (A, F, GH, K, MO, NY, S, US). BRAZIL. Acre: Cruzeiro
do Sul, Rios Jurud and Moa, 8 km above Cachoeira Grande, 07° 30'S, 73° 30' W, 27 Apr

1971 (fr), G. Prance et al. P12555 (TAN, MG, NY). Amazonas: Bas in of eee Mpio.
Sdo Paulo de Olivenga, near Esperanga, Dec 1935 (fr), B. Krukoff 7063 (NY, U

—

Cybianthus kayapii may be confused with C. spricei (Hook. f.) G. eee
but may be recognized by the shorter petioles, heteromerous flowers and
mucronate leaf apices. However, the leaf blades with mucronulate apices
and subauriculate bases, and heteromerous flowers indicate Cybianthus kayapit
is more closely related to C. anthuriophyllus. From Cybianthus anthuriophyllus,
C. kayapii may be separated by the much smaller, flat, entire leaves with
entire margins, the deltate calyx lobes with entire margins, and the carnose
corolla.

21. Cybianthus anthuriophyllus Pipoly, sp. nov. (Fig. 17). Type: ECUA-
DOR. Napo: Cantén Gonzalo Pizarro, Rio Tigre, affluent of Rio Dear entering
from 73 km of Rd. from Lumbaque to El Reventador, 10 km S of Lumbaque, 00°
05'S, 77° 24' W, 900-1,100 m, 18-21 Feb | 1987 (pist. fl, fr), W. SR Neill
1584 (HoLorypeE: US; 1sorypes: K, MO, E).

Ob foli a oblanceol ata ab lamina ad Sala gre acacl contracto, a primo intuitu cum

t, sed: aD @a margini ib JU aminaribus if itegerrimis

ea Sane) inflorescentiis 28 (non 8— ae longis, pleiochasia cum eenaulie aon eris cymosis
(nec bipinnatips ei cum ramulis floriferis racemosis) lobulis calycinis subulatis (nec
ovatis) praeclare dista

Monoaxial treelet to 0.6 m tall. Stems terete, weakly woody, ca. 2 cm diam.,
sparsely glandular-papillate, glabrescent. Cataphylls ughtly pseudoverticillate,
coriaceous, linear-subulate, 4.0—6.5 cm long, 0.3—0.6 cm wide, densely and
prominently punctate-lineate, sparingly glandular-papillate, glabrescent.
Leaves pseudoverticillate, erect; blades coriaceous, narrowly lanceolate, (56—)
104—110cm long, (14.5—)17—22 cm wide, apically broadly rounded or rounded
to a minute abrupt submucronate acumen 2 mm long, the blade gradually
tapering to the petiole, almost obtusish basally, midrib slightly raised above,

90 SIDA 18(1)

prominently raised below, the secondary veins ca. 42 pairs, impressed above
prominently raised below, nitid and epunctate above, purple (when fresh),
inconspicuously punctate and papillate-puberulent below, the papillae erect,
rufous, the margin thin, opaque, inrolled, prominently straight-serrulate,
the teeth alternatively larger and smaller (ca. 1 mm and 0.5 mm long, re-
spectively); petiole deeply canaliculate, thick, ca. 1.5 cm long, 0.8 cm diam.,
sparingly glandular-papillate. Staminate inflorescence: a pyramidal aiyisold
panicle, ca. 16 cm long, 23 cm e, the branches pseduoracemose (sym-
podial), the rachis densely g eal ate, the peduncles longer below,
shorter above; inflorescence branch bracts linear-lanceolate, 7-9 mm long,
2—3 mm wide, apically attenuate- —— sparsely glandular-papillate, the
margin entire; floral bracts acerose, 2-3 mm long, 0.4—0.8 mm wide; pedicels
1.0—2.5 mm long, angular, thin, - longer pedicels in the lower portion of
the inflorescence, the shorter ones above, densely glandular-papillate, the
papillae persistent. Staminate flowers heteromerous, membranaceous, hyaline,
the calyx (5—)6-merous, the corolla 5-merous; calyx subcotyliform 3.7—4.7
mm long, 0.4—0.7 mm wide, apically acuminate-acicular, prominently keeled,
the keel thickened hyaline, epunctate, sparsely glandular-papillate except
densely so along the margin, the margin entire; corolla subrotate, 7.4—10
mm long, the tube 0.7—0.9 mm long, the lobes lanceolate-subulate, 6.3—9
mm long, 2.7—3.2 mm wide, the apically long-acuminate, highly reflexed,
sparsely glandular-papillate without, very sparsely glandular-granulose within
above the junction within tube the margins densely glandular-papillate,
entire; stamens 2.2—4.1 mm long, the staminal tube conspicuous, coria-
ceous, 1.4—1.8 mm long, glabrous, elobate, the apically free portions of the
filaments ventrally recurved, 0.8—0.9 mm long, flat, glabrous, the anthers
ovate, ().3—0.4 mm long and wide, apically and basally emarginate, dorsifixed
less than _ length, the connectives prominently black punctate ventrally
and dorsally; pistillode conic, vestigial-O.8 mm long or absent. Pesti/late
inflorescence a pleiochasium, with branches cymose (sympodial), the rachis
densely glandular-papillate, the peduncles 1—3 cm long, longer below, shorter
above; inflorescence bracts resembling cataphylls but smaller, 1.3—1.6 cm
long, 1.0—1.3 mm wide, conspicuously punctate-lineate; inflorescence branch
bracts linear-lanceolate, 1.2—1.5 mm long, 0.2—0.4 mm wide, apically at-
tenuate-acicular, sparsely glandular-papillate, che margin entire; floral bracts
acerose, longer than the pedicels, 0.5—1.0 cm long; pedicels angular, thin,
1.2—7.5 mm long, the longer pedicels in the lower portion of the inflorescence,
densely glandular-papillate, the papillae persistent. Prsti/late flowers as in
staminate but calyx 6-merous, (2.1—)3—4.1 mm long, the tube 0.3—0.7 mm
long, 0.5—1.0 mm wide, apically long-attenuate-acicular; corolla (from dried
remnants) subrotate, hyaline, 5.O—7.1 mm long, the tube 1.0—-1.3 mm long,
the lobes linear-subulate, 4.0—5.8 mm long, 1.1—1.5 mm wide at base, apically

Pipoty, Cybianthus in Ecuador and Peru 91

subulate, highly reflexed, twisted and distally recurved at anthesis; stami-
nodia 1.2-1.3 mm long, the staminodial tube 0.9-1.2 mm long bearing
well-developed lobes alternate with the apically free filaments, the filaments
0.3—1.1 mm long, flat, glabrous, the antherodes malformed, mostly con-
sisting of undifferentiated tissue surrounding prominent black punctations
or suborbicular and 0.1—0.5 mm long, 0.1—0.6 mm wide, always wider than
long, apically emarginate, dorsifixed slightly less than _ length, the connectives
prominently black punctate dorsally; pistil obturbinate, 2.5—3.0 mm long,
the ovary 1.0—1.5 mm long, 0.8—1.3 mm wide, densely papillate and prominently
black punctate, the style 1.3—1.7 mm long, densely glandular-papillate, the
stigma large, capitate, lobes, the lobes lacinate, each lobe to 0.4 mm long,
early caducous, the placenta widely conic, bearing 4 uniseriate, exposed ovules,
the ovules on the periphery of the placenta. Bisexual fruit pink, globose, 4—
6 mm long and diam., the punctation prominent, brown when fresh (teste
coll.), red or black upon drying, the exocarp thin.

Distribution —Amazonian (“Oriente”) Ecuador and adjacent Peru (Loreto),
160—300(—1,100) m elevation.

Ecology and conservation status. —Cybianthus anthuriophyllus grows in pri-
mary tropical wet forest and premontane wet forest on terra firme above the
high water contour. It is found in primary forest as well as in secondary, but
it is not known whether the plant is cultivated in secondary forest situa-
tions. At this time, the species can be considered locally common but not
threatened.

Etymology.—The specific epithet refers to the unique shape of the adult
leaves, held erect 72 vivo and reminiscent of A ntiourinm ¢ crassinervinm ne

ocd. 5 and uses:—Ecuador: “namakuk” (Achuar Jivaro); “ rf’ (dialect
ene “challuo panga” (Quichua). Peru: oe Fe hachia’
“mututpash,” “kurip” (Mayna Jivaro), “sierra panga” (Quichua). Leaves crushed
and used as a fish poison( W. Lewzs et al. 14051); inner stem is scraped and
an infusion given to dogs to drink to improve their hunting abilities and to
enhance their stamina (W. Lewis et al, 11153); stem is scraped and its juice
put into a dog’s nose to produce sneezing (W. Lewis et al. 10475), or to “make
it an attack dog against thieves” (Lew7s et al, 12855).

Pararypes. ECUADOR. Napo: Cuyabeno-Punta Arenilla; Sep te an: fl), E. Bravo
& P. Gémez 49 (QCA); Canton Orellana, Yasunf National Park, MORGR
project km 10, 00° 29'S, 76° 34' W, 250 m, 29 Jun 1994 (fr), N. ai 448 (MO, ne
Pastaza: Kapawi (Amuntajf), rio Pastaza; Village area, 02° 31'S, 76° 4 ae 235 m,

29 Jul 1989 (ster.), W. Lewis et al. 14051 (MO). PERU. Bonet: Prov. ane
Puranchim, Rio Sinchiyacu, 02° 50'S, 76° 55' W, 200 m, 3-7 Dec 1988 oo W. Leu
et al, 14390 (MO); Washintsa and vicinity, Rfo Huasaqa, bee 20'S, 76° 20' W, 185 m,
16-26 Jun 1986 (ster.), W. Lewis et al. 11153 (MQ); Prov. Loreto: Nueva ae alem and
vicinity, Rfo Macusari, 02° 55'S, 76° 15' W, 220-300 m, 29 Dec 1985-3 Jan 1986 (fr),
W. Lewis et al. 10475 (MO); Pampa Hermosa and vicinity, Rfo Corrientes, 1 km S of junc-

92 Sipa 18 (1)

tion with Rfo Mucusari, 03° 15'S, 75° 50' W, 160 m, oe Dec ey alee fl), W. Lewes
et al, 10340 (MO); Vista Alegre, Rio Tigre, 02° 40! S, 35' W, 2 , 17 Mar 1987
(ster.), W. Lewrs et al, 12853 (MC

Cis anthuriophyllus is unique within subgenus Comomyrsine by its

small, marginal pectinate leaf serrations and pleiochasial inflorescence, ap-
pearing paniculate, but with sympodial primary and secondary branches,
and thus, cymose. The leaves appear subsessile, with the blade gradually
tapering to the deeply canaliculate petiole, a feature found otherwise only
in its closest congener, Cybianthus kayapii. However, Cybianthus anthuriphyllus
is Clearly distinct from C. kayapii because of the inrolled, pectinate-serru-
late leaf blade margin, the much longer, pleiochasial inflorescence, and subulate
calyx lobes. The extremely long cataphylls are the best developed in the
subgenus.

V. Pra iran subgenus Triadophora (Mez) G. Agostini, Acta Biol. Venez.

10:164. 1980. Wéeigeltia subgenus Triadophora Mez in Engl., Planzenr. [V. 236(Heft

9):291. 1902. Tyre Species. Wergeltia schlimit (Hook. f.) Mez in Engl. = Cybianthus
schlimit (Hook. f.) G. Agostini.

Correlliana D'Arcy, Ann. Missouri Bot. Gard 60:442. 1973. Type Specirs. Correlliana
spectabilis (Standl.) D'Arcy = Cybranthus schlimti (Hooker f.) G. Agostini.

As here interpreted, Cybianthus subgenus Triadophora is monotypic. Its

only species, C. schlimii, is easily recognized by its monoaxial habit and

autapomorphic rufous glandular comentum of malpigiaceous trichomes, leaf

blades with subepidermal fibers and pseudocataphylls (here defined as petiolate
cataphylls). The first full description of Cybzanthus schlimii is provided, along
with complete synonymy and complete exsiccatae for Ecuador and Peru,
and representative ones for other areas.

22. Cybianthus schlimit oe f.) G. Agostini (Fig. 3D, 6K, 6L), se
Biol. Venez. 10:165. 1980. Comomyrsine schlimii Hook. f. in Benth. & Hoo
Gen. Pl. 2: 644, 1 1876. Weigeltia aN .) Mez in Engl., Pflanzenr. IV. are
9):291. 1902. Correlliana schlimii (Hook. f.) D'Arcy, Ann. Missouri Bot. Gard 60:44
1973. Type: COLOMBIA. Mera: ea de se ae 300 m, Jan 1856 (stam. A,
J. Triana 7594 (u0LoryPe: K; tsoryees: COL, )

Weigeltia multiflora A.C. Smith., Bull. Torrey Bot. Club 60:387. 1933. syn. nov. Correlliana multi-
flora (A.C. $m.) D'Arcy, Ann. Missouri Bot. Gard 60:445. 1973. (A.C. Sm.) G. Agostini,
Acta Biol. Venez. 10:165. 1980. Type: BRAZIL. Maro Grosso: Near Tabajara, upper

o Machado, 23 Nov 1931 (stam. fl), B. A. Krukoff 1388 (HoLotypr: NY; isoryee: A).

ee spectabilis Scandl., Publ. Field Mus. Nat. Hist. Bot. Ser. 18:893. 1938. syn. nov.

Weiveltia spectabilis (Standl.) Lundell, Wrightia 4:169. 1971. Cian spectabilis (Standl.)
D’

4

nn. Missouri Bot. Gard. 60:443. 1973. Cybianthus spectabilis (Standl.) G.
Agostini, Acta Biol. Venez. 10:165. 1980. Type: COSTA RICA. ALaAjuELA: Cataratas
(Los Angeles) de San Ram6n, Apr 1935 (stam. fl), A. Brenes 20530 (HOLOTYPE: F; ISOTYPES:

CR, F).
Weiseltie triandra Aspl., Bot. Not. 1939:802. 1939. Type: COLOMBIA. Cauca: Near
Distrito El Tambo, 900 m, 31 Jul 1936 (stam. fl), K. von Sneidern 919 (HOLOTYPE: S).

Pipoty, Cybianthus in Ecuador and Peru ee)
Weigeltia schlimii (Hook. f.) Mez var. intermedia Moldenke, Phytologia 2:242. 1947. Type:
COLOMBIA. VAaL_e DEL Cauca: Pacific coast, Rfo Cajambre, San Isidro, 5-100 m, 2—

5 May 1944 (stam. fl), J. ae ecasas 17312 (HOLOTYPE: NY; 1soTyPEs: COL, F-2 sheets).
Monoaxial tree to 5 m. Stem terete, 0.8—2.0 cm diam., the wood dense,
minutely rufous glandular appressed tomentose, the trichomes malpighiaceous,
early glabrescent. Psewdocataphylls produced only irregularly, chartaceous,
subulate, ca. 2.0—3.0 cm long, 0.5—1.0 cm wide, apically acute, mucronate,
densely rufous puberulent, black lineate-punctate, the margin entire; peti-
ole subobsolete, to 0.2 cm long. Leaves tightly pseudoverticillate; blades
chartaceous, elliptic, oblong or oblanceolate, 25-65 cm long, 5.5—20 cm
wide, apically acute or subacuminate, mucronulate, the mucro often sclerified,
the acumen 0.5—3.0 cm long, base long-attenuate, the blade decurrent on
the upper portion of the petiole, midrib slightly elevated above, prominent
below, the secondary veins 9-13 pairs, prominent, the marginal veins loop
connected, conspicuously striolate by subepidermal fibers, these visible above
and below, sparsely rufous puberulent above, moderately puberulent be-
low, at times glabrescent, hydropotes absent, sparsely to densely punctate
or lineate-punctate below, the margin opaque, irregular, entire to roughly
serrate; petioles canaliculate, 1.0—3.0(—-10) cm long, 0.5—1.0 cm diam., abruptly
swollen basally, puberulent, glabrescent. Staminate Inflorescence a pinnate or
bipinnate columnar panicle 13—40 cm long, 3—20 cm wide, the rachis densely
glandular-papillate and rufous puberulent, the flowering branches racemose;
peduncle 8-15 cm long; inflorescence bract chartaceous, ovate, 9-15 mm
long, 2.4-4.5 mm wide, apically acute, densely rufous glandular puberu-
lent, conspicuously black punctate and lineate-punctate, the margin opaque,
entire; inflorescence branch bracts membranaceous, linear, 10-13 mm long,
1.9—2.1 mm wide, apically narrowly acute, mucronulate, minutely rufous
puberulent, orange furfuraceous lepidote, densely and conspicuously black
lineate-punctate, the margin opaque, entire; floral bracts membranaceous,
subulate, 0.8—1.3 mm long, 0.2—0.4 mm wide, sparsely rufous puberulent,
the margin entire; pedicel terete, 2.0-4.5 mm long, prominently black punctate,
densely papillate and rufous puberulent. Staminate flowers 3(—4)-merous, light
purple, then dull yellow; calyx chartaceous, cupuliform, 0.9—1.8 mm long,
the tube 0.3 mm long, the lobes subdeltate, 0.7—-1.6 mm long, 0.9-1.2
mm wide, apically acute, densely rufous puberulent, glabrescent, densely
and prominently black punctate, the margin flat, wide, SyeMats a ciliolate,
the cilia often caducous; corolla rotate, chartaceous, 3.0—5.0 mm long, the
tube hyaline, 0.8—1.0 mm long, the lobes elliptic or ae 2.24.0 mm
long, 1.6-2.2 mm wide, apically obtuse to rounded, subcucullate, invo-
lute, distally recurved 180° relative to tube, sparsely rufous puberulent without,
very sparsely glandular-granulose within basally, often glabrescent, very densely
and prominently black punctate except margin hyaline, irregular, glabrous,

94 Sipa 18(1)

entire; stamens 2.4—3.5 mm long, the tube membranaceous, inconspicu-
ous, 0.8—1.0 mm long, hyaline, epunctate, glabrous, elobate, the apically
free portions of the filaments 1.8—2.3 mm long, sparsely or epunctate, sparsely
rufous puberulent at first, glabrescent, the connective prominently punc-
tate, or inconspicuously so, the punctation orange, red or black, the anthers
cordate, 0.5—0.6 mm long, 0.7—1.0 mm wide, apically subacute to rounded,
base deeply cordate, dorsifixed at point less than 1/5 distance from apex;
pistillode none or conic, to 1.0 mm long, densely and prominently black
punctate and rufous papillate. Pisti//ate inflorescence as in staminate but more
columnar, (3.5—)8.0-18.5 cm long, (2.0—)3.5—6.0 cm wide, the branches
subcorymbose to rarely racemose; peduncle 1.5—6.8 cm long; inflorescence
bract ovate to elliptic, 4.5-12 mm long, 1.9—4.0 mm wide, conspicuously
black punctate; inflorescence branch bracts 5-15 mm long, 0.5—2.5 mm
wide; floral bracts 0.9-2.0 mm long, 0.3-0.6 mm wide; pedicels terete,
2.0—4.0 mm long, accrescent to 5.0 mm long and incrassate to 2.0 mm
diam. in fruit. Prste/late flowers as in staminate but 3-merous, calyx purple,
corolla yellow; calyx subcotyliform, 1.8-2.0 mm long, the tube 0.2—0.3
mm long, the lobes subdeltate to widely ovate, 1.6-1.8 mm long, 1.6—2.2
mm wide, apically acute, the margin slightly erose, conspicuously long glandular-
ciliate, the cilia often caducous; corolla 4.2-5.0 mm long, the tube 0.3—0.5
mm long, the lobes oblong, 3.2—4.5 mm long, 2.3—2.6 mm wide, apically
obtuse, somewhat cucullate, reflexed ca. 45° from tube axis, densely pu-
berulent without, sparingly glandular-granulose basally within; staminodes
to 2.5 mm long, the tube membranaceous, inconspicuous, to 0.5 mm long,
the filaments adnate to corolla lobe ca. 0.5 mm long, then apically free to
1.0 mm long, thick, terete, rarely punctate, the antherodes cordate, ca. 0.6
mm long, 0.7—0.8 mm wide, apically acute to apiculate, base widely cor-
date, the connective punctate, prominently or not; pistil clavate to obnapiform,
4,0—5.2 mm long, the ovary 1.8 mm long, 0.5—-1.6 mm wide, prominently
black punctate and rufous puberulent, the style to 1.3 mm long, densely
rufous puberulent, the stigma capitate to lobed, 3(—-4)-lobed, the placenta
deeply cupuliform, bearing 2 ovules exposed apically. Fruit globose, 1.0—
1.3 cm long, 1.0—2.0 cm diam., at maturity, the exocarp bright orange to
red-orange, prominently black punctate.

Distribution. —From the Atlantic Slope (Dpto. Rio San Juan) Nicaragua,
to state of Pando, Bolivia and adjacent Amazonia of Brazil, 0—1,8¢

Ecology and conservation status. —Cybianthus schlimti occurs in a variety of
habitats, from lowland to premontane tropical moist, wet and pluvial for-
est. It is locally common, but restricted to primary forest. Therefore, it should
be considered threatened.

Etymology.—The species is named for Louis Joseph Schlim, a Belgian plant
collector working for J.J. Linden in Brussels, who collected extensively around

Pipoty, Cybianthus in Ecuador and Peru 95

Caracas to the Venezuelan Andes, and from there, to the grasslands of Meta
and the Sabana de Santafé de Bogota, Colombia, during the period 1841-
1852. Schlim also collected with Nicolas Funck later in Venezuela.

Local names and uses.—Peru: “napi tsuake” (Huambisa).

Representative specimens examined. NICARAGUA. Rio San Juan: El Relos, midpoint
between E] Castillo and Delta de San Juan, 0-50 m, 23 Mar 1961 (stam. fl), G. Bunting &
L, Licht 775 (EF, NY). COSTA RICA. Alajuela: 15 km NW of Arenal by air, 2 km NW of
Nuevo Arenal on Rd. to Tilardn, then 3 km NE on Rd. to San Rafael de Guatuso, then 2
km W on Rd. to Finca Cote, 10° 34' N, 84°54' W, 700 m, SE side of Lago Cote, 30 Apr
1983 (fr), R. Liesner et al. 15093 (CR, MO, WIS). Heredia: Zona Protectora, N slopes of
Volcan Barba. betweem Rio Peje and Rfo Guacimo, along Quebrada Cantarana, 300—400
m, 18 Jan 1983 (stam. fl), M. Grayum & G. Shatz 3170 (CR, DUKE, MO); Finca La Selva,
Rio Puerto Viejo 2 km E of jct with Rfo Sarapaquf, 10° 26' N, 84°00! W, 100 m, 14-17
Jun 1968 (stam. fl), Wo Burger & R. Stolze 5803 (CR, F, MO, NY). Limon: Near Finca
Castilla, 30 m, 24 July 1936 (ster), C. Dodge & V. Goerger 9283 (F, MO). San José: Estacion
Carrillo, Cafion del Rio Sucio, 450-700 m, 12 Nov 1983 (pist. fl), 1. Chacén G G. Herrera
1720 (CAS, CR, MO, NY). PANAMA. Bocas del Toro: Cerro Bonyic, above Quebrada
Huron, 180-400 m, 13 Apr 1968 (fr), J. Kirkbride G J. Duke 610 (MO). Colon: Base of
Cerro Bruja, along Rio Escandaloso, above Mina Boquerén, No. 2, 47.5 km from Tran-
sisthmian Hwy on Rd. to Salamanca, 09° 50'N, 79° 32 ‘W, 10-200 m, 18 Mar 1982 (stam.
Al), S. Knapp & W. J. Kress 4282 (MO, NY, PMA). Darién: Rio Tuquesa, Tuquesa Mining
Co. camp, Charco Peje, 250 m, 7 Jul 1975 (stam. fl), 8. Mort 7015 (MO, SCZ). Panama
Cerro Campana, 800 m, 22 Jun 1967 (fr), T: Croat 17167 (MO). San Blas: Trail from Rio
Estadi to Cerro Banega, 300—530 m, 09° 23’ N, 78° 51' W, 21 Dec 1985 (stam. fl), G. de
Nevers & H. Herrera 6642 (CAS, MO, PMA). COLOMBIA. Antioquia: 6 km E of Guapa,
53 km S of Turbo, 240 m, 13 May 1945 (stam. fl), 0. Haught 4660 (US); Mpio. Anori,
Corregimiento de Providencia, Buenos Aires, 500-600 m, 4 Feb 1972 (fr), D. Soejarto 3205
(HUA, MO, NY); Vicinity Planta Providencia, 26 km 8, 23 km W (by air) of Zaragoza,
07° 13' N, 75° 03' W, valley of Rio Anorf between Dos Bocas and Anori, 1 Jun 1976 (fr),
J. Shepard 323 (COL, WIS); Vicinity Medellin, 20 Aug 1927 (stam. fl), R. Toro 356 (MEDEL,
NY); Medellin-Bogota Hwy, sector Rio Samana-R {fo Claro-San Luis, 400-1,000 m, 24 Aug
1982 (fr), J. Herndndez & S. Hoyos 483 (COL, HUA); Mpio. Urrao, Boundary of Parque
Nacional Natural Las Orquideas, Vereda Calles, Permanent Inventory, Premontane Rainforest,
left bank of Rio Calles, 06° 32' N, 76° 19' W, 1,450-1,500 m, 30 Nov 1993 (ster.), J.
Pipoly et al. 17406 (BRIT, JAUM, MO). Boyacé: Region of Cerro Chapén, extreme W
part of Boyacd, NW of Bogoté, 2,300 m, 31 Jul 1932 (fr), A. Lawrence 370 (A, NY, S); El
Umbo region, 1,000 m, 12 Oct 1932 (stam. fl), A. Lawrence 530 (A, BM, F, G, GH, MO,
NY, 8, U, UC, US). Chocé: Mpio. Quibdé, Quebrada La Platina, Hwy to Medellin, 25 Sep
1983 (fr), L. Arias et al. 134 (MO); Mpio. San José del Palmar, along Rio Torito (affluent of
Rio Habita), W slopes, 850-950 m, 15 Mar 1980 (fr), E. Forero et al. 7350 , 16 Mar 1980
(fl bud), E. Forero et al. 7393 (COL, MO), Vereda Portachuelo, Hacienda Barro Blanco, 1,350
m, 15 Jan 1983 (fr), P. Franco et al. 1325 (COL); Rio Mecana, ca. 10 km E of Mecana,
15'N, 77° 25' W, 100 m, 7 Mar 1983 (stam. fl), A. Gentry G A. Juncosa 41072 (COL,
JAUM), 710-880 m, 8 Jan 1984 (stam. fl), A. Juncosa ee (COL, MO, JAUM); ae
Novita, vereda Curundo, left bank, Rio Ingara, 550 m, 1 Dec 1983 (fr), BP. Franco et al.
1059 (COL); Rio Nuquf, 400 m, 25 Jan 1947 (stam. fl), O. Haught 5479 (COL, US); La
Mojarra, upriver from Istmina, 05° 12' N, 76°37' W, 30-60 m, 5 Nov 1983 (fr), A. Juncosa
1255 (COL, JAUM, MO, NY); S of Rio Condoto, between Quebrada Guarapo and Mandinga,
120-180 m, 22-28 Apr 1939 (fr), E. Killip 35675 (COL, US). Cundinamarca: Cordillera

96 Sipa 18(1)

Bogota, 2,000 m, Sep 1855 (fr), J. Triana 4 (BM). Meta: Near Salitre, 6 Jan 1876 (stam.
fl), E. André 1151, 7 Jan Here (ster), E. André s.n. (K); Cafio Tigre, between Cafio Aguas
vee and Cafio Grande, 4.5 km SW of Villavicencio, 04° 07' N, 73°39! W, 500-550 m,
24 1943 (pist. fl), FR. ce 20148 (COL, US); Llano de San Martin, (stam. fl, pist.
fl mix nN H. Karsten s.n. (LE-2 sheets). Narifio: Mpio. Barbacoas, Corregimiento Ortiz y
oe hie Vereda El Barro, ae ania Rio Nambjf, ca. 5 km W de sy lai faldas
ccidentales de la Cordillera Occidental, 01° 18' N, 78° 08' W, 1,350—1,400 m, 1S
io oe fl), J. Pipoly, A. Cogollo, et al. 21095 (BRIT, FMB, JAUM, ca ce i
,450-1,500 m, 2 Sep 1997 (fr), J. Pipoly, A. ees et al. 21483 1250-1,350 m, 8 Dec
ae (fr), J. Pipoly, A. Cogollo et al. 21598 (BRIT, JAUM, PSO). Norte de Santander:
Ocafia, 1200 m, May 1846-52 (stam. fl), L. ie 686 (BR- 3 sheets, F-2 sheets, G-BOIS-
2 pacar G-DEL, MA-2 sheets). Santander: 8 km SE of Barrancabermeja, Rio one 200
, 28 Aug 1954 (stam. fl), R. Castatieda 4746 (COL), vicinity Barrancabermeja, ~_
Rios Sogamoso and Colorado, 100— eee m, : of Rfo Sogamoso, Camp Mesa, 8 Jan 1935
(stam. fl), O. Haught 1502 (A, COL, FE N JS). Valle del Cauca: Cordillera en ty
W slope, along Rio ye ae left side, - ae 1,250—-1,400 m, 13 Dec 1943 (stam.
fl), J. Cuatrecasas 15474 (VALLE s Bahfa Malaga, Quebrada Algeria, new Rd. to military
base,O 4° 02'N, aa 22 W, 50 m, A. Gentry et al. 53326 (COL, MO, US); Bank se Rio
Digua, Rio San Juan, below OMe to the right of river between km 52 and 53, 1300—
1500 m, 19 Mar 1947 (fr), J. Cuatrecasas 23855 (COL, F-3 sheets); Calima ia. ae
1700 m, 17 Sep 1966 (stam.) S. ee 2109 (MO, VALLE). Without locality, date, (stam.
fl), J. Matis 449 (MA), 2919 (MA), 3907, 5102a¢ (MA, US), Warscewicz s.n. a destr., Photo-
F Neg. 4858). ECUADOR. Carchi: ay Bes plain above oO Donoso and Rio eee
01° LO' N, 78° 18—31' W, 280-450 m, 18 Feb 1984 (sta ee ep (MO), S
Trail, Gualpi Chicé Area of Awa Reser ve, near encampment, er N, 78° 16' W, 1,33 6
m, 19 Jan 1988 (stam. fl), W Hoover et al. 2815 (MO, QCNE), Gualpi Medio Community,
Awa Reserve, 900 m, 21 May 1992 (bud), C. Quelal et al. 764 (MO, QCNE) El Pailon, 45
km below arene ba path to Tobar Donoso, 800 m, 1 Dec 1979 (fr), M. Madison &
.. Besse 7201 (AAU, F, QCNE, SEL); Cantén Tulcain, Reserva Indigena Awa, Parroquia
Tobar Donoso, sector El Baboso, 00° 53! - 78° 20' W, 1,600 m, 3 Oct 1991 (A, fr), G.
Tipaz et al. 260 (BRIT, MO, QCNE), (fr), 377 (BRIT, MO, QCNE), Centro El Baboso, 00°
53' N, 78° 25' W, 1,800 m, 17-27 Aug 1992 (fr), G. Tipaz et al. 1950 (BRIT, EF MO,
CNB). Panes Cantén San Lorenzo, Reserva Indigena Awa, Cafion del Rio Mira,
10 km W of Alto Tambo, Comunidad “La Unién,” 01° 02' N, 78° 26' W, 250 m, 16—26
Mar 1991 (fr), D. Rubio et al. 1262 (MO, QCNE). Sucumbios: Cantén Cl Agrio, Reserva
Cuyabeno, Laguna Grande, Near NEOTROPIC eee OOS00' S76" LI Ww. 230 mm, 15
Nov 1991 (stam. fl), W. Palacios et al. 9269 (BRIT, MO, ee ). Zamora-Chinchipe:
Cancon Nangaritza, Valle del Rio Nangaritza, san 04° 18'S, 78° 40' W, 1,200 m, 10
ne nee (pist. . W. Palacios 6734 (BRIT, MO, QCNE), a military camp, 04° 16!
5 W, 97( 20 Oct 1991 (stam. fl), W Palacios et al. 8483 (BRIT, MO, QCNE).
mn Mtn tere Cia mino de chi ings entsa, 180-250 m, 2 May 1973 (stam. fl), R.
any 728 (MO), Valle del Rio Santiago, 03° 50'S, 77° 40' W, Quebrada Caterpiza, 2—3
km behind Caterpiza, 200 m, 4 Jan ees (stam. fl), S. Tungui 549 (MO). Loreto: Prov.
Maynas, Guarnicién Pijuayal, near Pebas, 130 m, 7 Sep 1988 9stam. fl), C. Diaz et al. 566
(MO). Ucayali: Prov. Padre ee Boquer6én del Padre Abad, 400 m, 20 May 1969 (fr), J.
Schunke 3068 (F-2 sheets, US, USM). BRAZIL. Amazonas: Mpio. Humayta, on plateau
between Rio Livramento & a Seer 7-18 Nov 1934 (stam. fl), B. Krukoff 7290 (A,
NY, S, U); Sao Paulo de Olivenga, near Esperanga, Dec 1935 (ster.), B. Krukoff 7603 (K,
NY), Behind Sao Paulo de Olivenga, 16 Aug 1973 (fr), E. Léeras et al. P17315 (GB, INPA,
MG, NY); km 500, Manaus-Humayta Rd., 17 Sep 1980 (stam. fl), S. Lowrie et al. 52 (INPA,

Os

‘cal

Prpoty, Cybianthus in Ecuador and Peru 97
MG, NY). Mato Grosso: Near Tabajara, upper Rio Machado, Nov—Dec 1931 (stam. fl), B.
Krukoff 1377 (A, NY). Rondénia: Sao Lourengo, cassiterite mine, 20 km N of S. Lourengo
on Rd. to “A Macisa” Mine, 15 Jul 1979 (stam. fl), C. Calderén et al, 2852 (INPA, US).
BOLIVIA. Pando: W bank of Rio Madeira, 3 km above Abuna, 13 Nov 1968 (stam. fl),
G. Prance et al, 8388 (LPB, MG, INPA, NY).

The “pseudocataphylls” referred to in the description are poorly devel-
oped and do not occur ina regular phyllotactic spiral as is found in subge-
nus Comomyrsine. Likewise, their morphology is essentially that of a leaf arrested
at different stages of development, and as such, do not have a distinctive
morphology.

Northern populations from Nicaragua and Costa Rica corresponding to
the type of C. spectabilis oS Ardista spectabilis) differ in their smaller flowers
and anther connect eglandular, a feature which is more a function
of ecotype then anything else. Eglandular anthers and entire leaves may be
found in very lowland wet forest populations in the Darién of Panama, and
the Chocé of Colombia.

The type of Wergeltia multiflora A.C. Sm. represents populations whose
inflorescences are less branched (although the duplicates of the type collec-
tion vary in that regard) and the largest flowers of the species. They, like
many of the Panamanian populations, have entire leaves and are otherwise
inseparable from the type of Weigeltia triandra Asplund.

The collections of Kayap from Amazonian Peru are referred to this spe-
cies, despite their longer petioles.

VI. ivan subgenus Nene (A. DC.) G. Agostini, Acta Biol. Venez.
56. 1980. ay A. DC., Trans. Linn. Soc. London, Bot. 17:102. 1834.
aie sect. Weigeltia (A. DC.) Miq. in Mart., Fl. Bras. 10:299. 1856. Type SpE-
cies: Salvadora surnamensis Spreng. Tent 7. 1828. =Cybianthus surinamensis (Spreng.)
G. Agostini.

Polyaxial dioecious or rarely, monoecious shrubs or small trees. Roots posi-
tively geotropic. Trwnk distinguishable, growth dynamics following Rauh’'s
Architectural Model (in ours), rarely Corner’s Model (not in Ecuador or Peru)
sensu Hallé et al. (1978). Bark grey to beige, thick, vertically fissured. Branchlets
sessile furfuraceous lepidote or rarely, rufous tomentose, often glabrescent
Cataphylls and pseudocataphylls absent. Leaves alternate, rarely subopposite,
with minute sessile rufous furfuraceous lepidote scales abaxially. Inflorescence
a pinnate or bipinnate panicle, rarely a simple raceme; peduncle 0.5—2 cm
long. Flowers 4- rarely 5-merous; calyx valvate, crenate or rarely entire, gla-
brous or rarely glandular-ciliolate, punctations red or black, prominent or
not; corolla rotate to subrotate, the lobes imbricate, glandular-granulose
only at the junction with the corolla tube, the punctations red or black,
conspicuous, or rarely, prominent; stamens and staminodes united to form
an inconspicuous or conspicuous tube, the tube without lobes alternating

98 Sipa 18(1)

with the apically free filaments, the filaments one to three times longer than
the anthers, terete, and recurved proximally, the anthers subglobose, or widely
ovoid, versatile, apically acute to emarginate, basally widely cordate, de-
hiscent by wide longitudinal slits, the connectives mostly prominently red
or black punctate; pistil conic, pyriform or obturbinate, the ovary translu-
cent glandular-lepidote or glabrous, the style capitate-lobate, the lobes entire;
pistillode minutely conic, or at times, absent. Frit globose or depressed-
globose.

Cybianthus subgenus Weigeltia contains approximately 46 species in South
America and the Caribbean. Five species are known from Ecuador and Peru;
they are restricted to the lowlands and premontane forests on the lower slopes
of the Western Cordillera in Ecuador and the Eastern Cordillera in Ecuador
and Peru.

KEY TO SPECIES OF CYBIANTHUS SUBGENUS WEIGELTIA

. Inflorescence bipinnately paniculate; petioles canaliculate; staminate flowers

with stamens shorter than corolla.

2. Branchlets terete; leaf blades apically long attenuate-subulate, basally long-
attenuate.

3. Branchlets 2.5—4 mm diam., minutely rufous-lepidote; leaves
pseudoverticillate, the blades membranaceous, prominently black punctate-
lineate below, margins subentire to — serrate; petioles 0. 5
cm long; calyx membranaceous, 1—1.3 mm long, the lobes obtuse, promi

ently punctate, glandular-ciliate on BIG TAPS he nessaastsheaeones 23. C. poeppigit
en ets 5.5—6 mm diam., minutely Ferrugineous tomentellous;
ae alternate, the blades eraeaecain. minutely and prominently

puncticulose below, margins entire; petioles 2-3. 5 cm long; eae
chartaceous, 0.7—1 mm long, the lobes acuminate, epunctate, glabrot
along the margin. 24. C, pee isaeiets

bo

Branchlets angulate or winged; leaf blades apically acute or es -acumi
nate, basally acute or cuneate.
Branchlets angulate, 8— 10mm diam. conspicuously rubiginous eae
lineate; leaf blad ispicuously rubiginous punctate-lineate
below; petioles 2.5—3 cm lc anes staminate calyx ee ea 1.4—
1.6 mm ee the lobes apically acute; staminate corolla membrana-
ceous, 2.3—2.5 mm long; stamens 1.8—2 mm long. ................... 25. C. vasquezit
oo . winged, (2.5—3—)5—6 mm diam., epunctate; leaf blades
chartaceous, inconspicuously pellucid punctate; petioles 2—2.5 cm long;

staminate calyx chartaceous, 1—1.2 mm long, the lobes apically
aa ar staminate corolla chartaceous, 1.2—1.3 mm lo ny sta-
—l. 1 long 26. C. cenepensis
. Inflorescence racemo ae marginate; staminate flowers with stamens
exserted or saline corolla. 27. C. nanayensis

23: Seats seed Ye Mez in Engl., Pflanzenr. IV. 236(Heft 9):218.
1902. Type: PERU. Lorero: Prov. Maynas. Tocache, without elevation, without
date, (stam. fl), F. are s,m. (HOLOTYPE: W),.

PipoLy, Cybianthus in Ecuador and Peru 99

Mee albiflora A. C. 8m., Amer. J. Bot. 27:546. 1940, syn. nov. Sateen albiflorus
m.)G. Agostini, Acta Biol. Venez. 10:157. 1980. Type: COL ;
ee 70-110 m, 20-30 Apr 1939 (stam. fl), E. Killip 35372 oe NY;
IsoTyPEs: A, BM, US-2 sheets).
Cybianthus ey Lundell, Wrightia 5:195. 1975, syn. nov. Type: COLOMBIA. Cuoco:
erro Mal{, on border with Panama, 1,200—1,400 m, 17 Jan 1975 (stam. fl), A. Gen-
try G&S. oe 13709 (HOLOTYPE: LL; IsorypPE: MQ).

Shrub or small tree to 4(—7) m tall. Branchlets thin, terete, 2.5—3.5(—4) mm
diam., minutely rufous-lepidote. Leaves in loose pseudoverticels; blades
membranaceous, elliptic, oblanceolate, lanceolate or rarely obovate, (6—)6.8—
18.5(—24) cm long, (2.2—)3.5—6.5(—8.5) cm wide, apically acuminate to
subacuminate-attenuate, basally cuneate to acute, not decurrent on the petiole,
midrib depressed above, prominently raised below, the secondary veins 7—
12(-14) pairs, rufous lepidote above and below early glabrescent above, tardily
glabrescent below, sparsely black punctate and densely black punctate-lineate
below, the margin flat, subentire to obtusely serrate; peaores cana Cue
0.5—1.5 mm long, minutely rufous-lepidote
monomorphic, bipinnately paniculate, somewhat oe soiierines
malformed and appearing racemose, 8—15 cm long, 8-10 cm wide, the ra-
chis densely glandular-papillate, the lowers racemose; inflorescence bracts
unknown; floral bracts membranous, linear-lanceolate, 1—2.5(—3.5) mm long,
0.3—0.8(-1.2) mm wide, apically attenuate, densely glandular-papillate,
epunctate, the margin glandular-ciliate; pedicels cylindrical, (0.7—)1.6—1.9(—
2.5) mm long, densely glandular-papillate. Staminate flowers 4-merous, yel-
low or yellowish-green; calyx membranaceous, cotyliform, 0.8—1.3 mm long,
the tube 0.2—0.3 mm long, oe the lobes widely ovate to suborbicu-
lar, (0.6—)1 mm long, (0.5—)0.6—1 mm wide, apically obtuse, prominently
punctate, the margin hyaline, irregular, erose- Saaeey ee densely glandular-
ciliate; corolla membranaceous, rotate, (1.7—)1. .2(—2.6) mm long, the
tube (0.3—)0.4—0.6 mm long, glabrous without, pa granulose within,
the lobes suborbicular, (0.9—)1.2—1.6(—2) mm long, (0.8—)1.2—1.7(—2) mm
wide, apically obtuse, prominently punctate, glabrous without, glandular-
granulose medially at stamen base within, the margin hyaline, irregular,
erose; stamens (1—)1.4—1.6 mm long, the tube inconspicuous, hyaline, (0.3—)
0.4—0.6 mm long, densely glandlar-granulose within, the apically free por-
tions of the filaments (0.3—)0.4—0.5 mm long, the anthers ovate-triangu-
lar, 0.3—0.4 mm long, 0.2—0.3 mm wide, apically rounded, basally obtuse,
the connective epunctate; pistillode conic, 0.3—0.4 mm long, densely ru-
fous glandular-papillate. Pistil/ate flowers as in staminate, but calyx 1—-1.3
mm long, the tube 0.2—0.3 mm long, the lobes deltate to widely elliptic,
(.8—1 mm long and wide, the margin opaque; corolla 2—2.3 mm long, the
tube, 0.6-0.7 mm long, the lobes 1.4-1.6 mm long, 0.6—-0.7 mm wide,

100 Sipa 18(1)

glandular-granulose within above filament junction with tube, the margin
often erose; staminodes resembling stamens but |—1.3 mm long, the staminodial
tube ca. 0.6 mm long, the apically free portions of the filaments to 0.3 mm
long, the antherodes suborbicular, to 0.2 mm long and wide; pistil obnapiform,
1.2—1.3 mm long, the ovary 0.8—0.9 mm long, 0.3—0.5 mm wide, densely
translucent-lepidote, the style 0.3—0.4 mm long, the stigma bilobed, the
lobes decurrent, ca. 0.4 mm long. Fruit depressed-globose, 3.5—4.5 mm
long, 4.5—6 mm diam. when dried, red, then black at maturity, inconspicuously
punctate, exocarp thin.
Distribuiton.—Known from the easternmost Darién of Panama south to
Amazonian Peru and Brazil, 1OO—1,800 m
Ecology and conservation status.—Cybianthus poeppigii is a broad ranging
polymorphic ochlospecies, occurring in primary premontane wet and plu-
vial forests, from the transition zone with lowland forests, to the transition
zone to cloud forests. Recent fieldwork in the Cordillera Occidental of Co-
lombia has shown it is a conspicuous element of primary pluvial premontane
forests, with a density of approximately 20 individuals per hectare, clus-
tered mostly along the margin of the windward side of the forest, and along
streambanks above the high water level. While locally common, its restricted

primary habitat verifies its threatened status.

Etymology.—The specific epithet honors Eduard F. Poeppig (1798-1868),
professor at Leipzig, explorer and plant collector, who made numerous valuable
contributions to our knowledge of the Peruvian Amazon Basin.

Representative specimens examined. PANAMA. Darién: Serran{a del eet Panama/
Colombia frontier, Cerro Tacaracuna, Cerro Mali, summit, 1,400 m, 17 Jan | a
A. Gentry G&S. Mori 13665 (LL-TEX, MO-2 sheets), W ee C cr Trac 1,800—
1,850 m, 31 Jan 1975 (fr), A. Gentry GS. Mori 14023 (COL, LL-TEX, MO-2 ts); Serrania
de Pirré, Cerro Pirré, above Cana Gold Mine between Rios Cm and ae aa 1,000—
1,310 m, 27 - 1976 (stam. fl), T; Croat 37785 (LL-TEX, : , Ph , SW ridge
leading to Alturas de Nique, Panama/Colombia border, | ,100—1,200 m, 30 Dec 1980 (stam.
fl), Rk. Hartman " is 12461 ( ). COLOMBIA. Antioquia: Mpio. De Anori, Corregimiento
vlc Rio Anorr Valley, between Dos Bocas and Anorif, 400-900 m, 24-31 May
fr), D. Soejarto et al. 4090 (A, COL, F, HUA, MO); Mpio. Sons6én, Rio Verde ee
ee ‘La Soledad,” 1,430—1,800 m, o iaa 192 oe ” G. Gutierrez 1186 (F, MEDEL,
, UC); Mpio. San Carlos, Corregimiento Alto de {, Vereda Miraflores, on 900
m, 15 Jun 1989 (stam. fl), R. Fonneera et al. 3076 ee HUA). Mpio. Urrao, Parque
Nacional Natural Las Orquideas, Vereda Calles, Permanent ae, aoe bank, Rio Calles,
06° 32'N, 76° 19! W, 1,450 m, 27 Nov 1993 (ster.), J. Pipoly et al. 17183 (BRIT, JAUM,
MO), 1,450—1,500 m, 29 Nov 1993 (ster. seedling), J. Pipoly et oe 17360 (BRIT, JAU
MO). Choco: Mpto. San José del Palmar, vereda “El Tabor,” 1,540 m, 18 Jan 1983 (pist.
fl), PB. Franco et al, 1469 (COL). Vaupés: Rio Pacoa Sia of as Apapéri. 00° 20'N,
71° 20' W, 300 m, 7-12 Feb 1952 (pisc. f), R. Schultes G 1. Cabrera 15423 (COL, GH,
US). ECUADOR. Morona-Santiago: Macuma, 50 km oy Macas, 21 Mar 1 1973 (pist. fl),
H, Lugo 3633 (GB); Rio Cuyes and ee Rd., 03° 25'S, 78° 35' W, 800 m,
| Nov 1986 (stam. fl), Wo Palacios 1460 (MO, US). Napo: Between Tena and Napo. | Jan

Prroty, Cybianthus in Ecuador and Peru 101
1940 (stam. fl), E. Asplund 10209 (S), 7 Jan 1940 (stam. fl), E. Asplund 10302 (S), Tena,
400 m, 3 Apr 1935 (stam. fl), Y. Mexia 7206 (NY, UC, US); Reserva Biol6gica Jatun Sacha,
8 km from Puerto Misahuall6, right bank, Rio Napo, 01° 04'S, 77° 36' W, 450 m, C.
Cerén 2585 (MO, QCNE). Pastaza: Mera, 1,100 m, (stam. fl), E. Asplund 18717 (S), 1,500
m 29 Dec 1958 (stam. Al), G. Harling et al. 9764 (GB); Loracachi, on path to Ce
01° 38'S, 75° 58! W, 25 May 1980 (fr) J. Jaramillo et al. 30984 (AAU, QCA), 31 Ma
1980 (fr), 31579 (AAU, QCA). Tungurahua: Rio Negro, 1,200 m, 13 Mar 1980 (ne fl),
G. Harling & L. Andersson 17255 (GB), 10 km E of Paquisha, 1,400-1,500 m, 13 Apr
1985 (stam. fl), G. Harling & L. Andersson 24097 (GB). PERU. Amazonas: Rio Santiago,
3 km behind Caterpiza, 14 Nov 1979 (fr), V. Huashikat 1248 (MO, NY); Tingo Maria, 25
Jul 1940 (stam. fl), E. Asplund 12497 (S). Pasco: Prov. Oaxapampa, Cabeza de Mono, Palcazu
Valley, 10° 20'S, 75° 18' W, 320 m, 11 Jun 1983 (fr), A. Gentry et al. 41880 (MO); Ozuz
to Rio Lobo, 10° 19'S, 75° 16' W, 400-500 m, 10 May 1985 (fr), R. Foster & B. d’Achille
10013 (F). San Martin: Quebrada de eee (right margin Rfo Huallaga), Prov. Mariscal
Caceres, Dtto. Tochache Nuevo, 400-500 m, 1 Jul 1974 (stam. fl), J. Schunke aN (MO,
NY). BRAZIL. Amazonas: tes Dec 1900 (stam. fl), E. Ule 5160 (HBG).

Cybianthus poeppigit was mistakenly placed by Agostini (1980) in subge-
nus Cybianthus, but the versatile, dorsifixed anthers clearly place it in sub-
genus Weigeltia. | had reported earlier (Pipoly 1983a) that Cybianthus albiflorus,
its synonym, was closely related to C. dawrencei Moldenke. However, the
bipinnate inflorescences with racemose flowers, stamens shorter than the
corolla and usually long petioles indicate that Cybianthus poeppigit is more
closey related to C. /ongifolins Miq., a vicariant species of the southwestern
Amazon Basin of Brazil and adjacent Bolivia. The glandular granules of
the staminal and staminodial tubes represent a unique (autapomorphic) character
state within the genus.

e holotype of Cybianthus poeppigii is staminate, as are those of its taxo-
nomic synonyms. The type of Wezgeltia albiflora (Cybianthus albiflorus) rep-
resents populations with entire, irregularly margined leaves, but is other-
wise qualitatively identical with that of C. poeppigi. Likewise, the type of
Cybianthus gentryi Lundell represents isolated montane populations of the
Darién/Chocé regions, and exhibits more notable lineate-punctations, much
smaller leaves, and abbreviated inflorescences. According to annotations by
Killip at US, description of another taxon was at one time contemplated,
based on the fact that the populations of this species from near Tena, Ecua-
dor, have roughly serrate leaf margins.

24. Cybianthus pseudolongifolius Pipoly, sp. nov. (Fig. 18). Type: PERU.
Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de Mono, 5—6 km W of Iscosacin
10° 12'S, 75° 14' W, 325 m, 13-19 Apr 1983 (fr), D. Smith 3808 (HoLotyPE: MO;
ISOTYPES: US, USM).

Quoad folia magna chartacea, inflorescentias bipinnatipanicul atas, | I dicellos cylindricos,
lobos calycinis ovatos, ca. 1/3 connatos, petiolos canaliculatos oe longifolio arcte accedens, sed
ab ea ramulis teretibus (non angulatis), 5.5—6 (non 3.5—4) mm diammetris, laminis anguste

oblanceolatis (non ellipticis vel lanceolatis) desuper sordidis hes nitidis) ad apices longi-

102 Sipa 18(1)

Fic. 18. Aa ia a tal a Pipoly. A. Habit, ae bipinnate panicles. B. Pe-

duncle and axillane leaf, showing canaliculate petiole. C. Detail of prominently puncticulose

abaxial leaf nae D. Pedicel, calyx and fruit, showing an and erose calyx lobes. A &
C, drawn from isotype; B & D, drawn from holotype. Figure drawn by Linda Ellis.

Pipoty, Cybianthus in Ecuador and Peru 103

attenuatis et subulatis (nec acutis vel acuminatis) ad bases longi-attenuatis (nec acutis),
icellis 2 8- ee 0. = Dr mm yous necnon

on

AAC esse Oras 8— 1S (non De “Oe ae ped

Gels to3m Sell Branchlets terete, 5.S—G6 mm diam. Aare and mi-
nutely ferrugineous tomentellous. Leaves alternate; pe chartaceous, nar-
rowly oblanceolate, (22—)26.5—33(—36.5) cm long, (5—)6.5—8.5 cm wide,
apically long attenuate, subulate, basally long-attenuate, decurrent on the
petiole, sordid and glabrous above, pallid, minutely and prominently pel-
lucid puncticulose and minutely ferrugineous puberulent below, midrib slightly
raised above, prominently raised below, the secondary veins 13—18 pairs,
the margin entire, glabrous, flat; petioles canaliculate, (2—)2.5—3(—3.5) cm
long, glabrous above, minutely ferrugineous puberulent below. Stamznate
inflorescence: unknown. Pistillate inflorescence: a lax bipinnate panicle, 8-13
cm long, 1.5—4 cm wide, densely ferrugineous puberulent, glabrescent; secondary
inflorescence bracts unknown; floral bracts unknown; pedicels cylindrical,

—3 mm long, densely ferrugineous puberulent; P7sti//ate flowers unknown;
fruiting calyx chartaceous, cotyliform, 0.7-1 mm long, the tube 0.3—0.4
mm long, the lobes ovate, 0.5—0.7 mm long, 0.5—0.6 mm wide, apically
acuminate, densely and prominently red punctate, the margin hyaline, erose,
epunctate, glabrous. Fruit depressed-globose, 3—4 mm long, 5-6 mm diam.,
inconspicuously pellucid punctate, minutely longitudinally costate.

Distribution.—Known only from the type.

Ecology and conservation status. —C ybianthus psendolongifolius appears to be
restricted to the lowland primary forest of the eastern Andean slopes. The
Oxapampa Province of Pasco is home to numerous Peruvian endemics, and
it would not be surprising if the species was of extremely limited distribu-
tion or endemic. With only one specimen known, no determination can be
made of its true conservation status.

Etymology.—The specific epithet refers to its general likeness to Cybianthus
longifolius Miq., a lowland black water river species from Amazonian Brazil
and Venezuela.

Cybianthus psendolongifolins is most closely related to C. longifolius, but
differs by its terete branchlets, 5.5—6 mm in diameter, narrowly oblanceolate
leaf blades that are sordid above, long-attenuate and SHO Ce apically and
long-attenuate basally, longer inflorescences, much longer p epressed-
globose, minutely costate fruits. The Oxapampa region of Pasco contains a
number of endemic species and disjunct taxa and as such, is one of the most
important underexplored areas in Peru.

25. Cybianthus vasquezii Pipoly, sp. nov. (Fig. 19). Tyr: PERU. Lorero:
Prov. Alto Amazonas, Capahuari Norte, 02° 45'S, 76° 25' W, 220 m, 7 Jun 1981
(stam. fl), R. Vasquez & N. Jaramillo 1993 (HoLoTYPE: MO; Isorypes: AMAZ, BRIT,
FE, NY, US, USM).

104 Sipa 18(1)

|

Fic. 19. Cybianthus vasquezti Pipoly. A. Habit, cea punctate-lineate branchlet.
B. Inflorescence branch, show wing racemose-glomerulate floral arrangement. C€ nc
rolla. D. Abaxial calyx lobe surface. E. Abax al leaf surface, showing minute furufuraceous
scales and prominent punctate-lineations. EF Branchlet apex. A—F, drawn by Linda Ellis,
from holotype.

Pipoty, Cybianthus in Ecuador and Peru 105

P ] Sigs ] oan ] lod ] ‘ I

<a
ad Be ses acuta, petiolos canaliculatos, infl i bipi i icul ramulos

inflorescentiares dense Siento vel slomerulacos lobos calycines grosse Erenetos C. pees

8 puncte
(non alterna), laminis oblanceolatis ( li of ‘is), (9—)10— 1315. 3) (nec ae 5-8) cm latis,
petiolis 2.5—3 (nec 1.5—2) cm longis, jaderescuaciis 12-25 (non 3-6) cm longis, corollae
lobis ovatis (non ellipticis) ad ae acute rocundatis (nec emarginatis) ad bases abrupte
constrictis (nec rectis), secus margines g (nec integerrimis) confeste separabilis.

Terrestrial dioecious tree to 8 m tall. Branchlets angulate, 8-10 mm diam.,
densely and minutely rubiginous furfuraceous-lepidote, conspicuously ru-
biginous punctate-lineate below. Leaves pseudoverticillate; blades coriaceous,
oblanceolate, (21—)26—34.5 cm long, (6.5—)9-12.2 cm wide, apically acute
to subacuminate, basally acute, decurrent on the petiole 4-7 mm, glabrous
above, very minutely rubiginous furfuraceous-lepidote below, the midrib
slightly raised above, prominently raised and rubiginous punctate-lineate
below, the secondary veins 7-10 pairs, slightly impressed above, promi-
nently raised below, the margin entire, flat; petioles canaliculate, 2.5—3 cm
long, swollen below to 0.5—0.7 cm diam. basally, rubiginous furfuraceous-
lepidote at first, early glabrescent, conspicuously rubiginous punctate-lineate.
Staminate inflorescence: a bipinnate panicle (12—)14-17.5(—25) cm long, 2—3
cm wide; peduncle (1—)2—3.5 cm long; secondary inflorescence bracts chartaceous,
linear, 2-3 mm long, 0.3—0.6 mm wide, apically attenuate, densely glan-
dular-papillate, the margin entire; branches racemose-glomerulate (0.5—)1—
1.5 cm long; floral bracts membranaceous, linear, 1.6—1.8 mm long, 0.3—
0.4 mm wide, apically attenuate, densely rubiginous puberulent; pedicels
cylindrical, 0.3-0.5 mm long, glabrescent. Staminate flowers 4-merous, pink;
calyx membranaceous, cotyliform, 1.4—1.6 mm long, the tube ca. 0.2 mm
long, the lobes ovate, 1.2-1.4 mm long, 1-1.1 mm wide, apically acute,
sparsely and inconspicuously orange punctate, glabrous, the margin coarsely
crenulate, glabrous; corolla membranaceous, subrotate, 2.3—2.5 mm long,
the tube 0.2—0.3 mm long, densely glandular-granulose, the lobes ovate,
2—2.3 mm long, 1.7—1.8 mm wide, apically acutely rounded, abruptly constricted
basally, inconspicuously orange punctate without, sparsely gladular-granulose
behind the base of the filaments, the margin hyaline, coarsely crenate, gla-
brous; stamens 1.8—2 mm long, the tube carnose, squarrose, 0.2—0.3 mm
long, the filaments terete, 1.1—1.2 mm long, slightly curved proximally,
the anthers ovate, 0.5—0.6 mm long, 0.6-0.7 mm wide, apically rounded,
basally cordate, the connectives inconspicuously orange punctate dorsally;
pistillode obturbinate, 0.5 mm long, 0.6 mm diam., the stigma truncate,
minutely lobed. Pzsti//ate inflorescence as in staminate but (3.5—)5.5—9.5 cm
long, the branches glomerulate. Pisti//ate flowers as in staminate but red,
except white on corolla lobe apices; calyx obconic, 1.3—1.7 mm long, the
tube 0.6—0.8 mm long, the lobes very widely ovate to suborbicular, 0.7—

106 Sipa 18(1)

1.2 mm long, 0.8—1.2 mm wide, apically broadly rounded, corolla 2.3—2.7
mm long, the tube 0.6—0.8 mm long, the lobes suborbicular, 1.5—1.7 mm
long, 1.5-1.8 mm wide, the margin irregular, hyaline, staminodial tube
1-1.3 mm long, the apically free portions of the filaments 0.4—0.5 mm long,
the antherodes 0.3—0.4 mm long and wide; ans obnapiform, 2.2—2.4 mm
long, 1.3—1.5 mm diam, the ovary 1.3—1.5 mm long, the style thick, Pe
1.3 mm long, the stigma capitate, 4-lobed, the lobes recurved, the placenta
deeply cupuliform, the ovules 2—3, imbedded. Frait unknown.

Distribution.—Known only from the Department of Loreto, in Alto Amazonas
and Loreto Provinces, Peru, at 160—220 m elevation.

Ecology and conservation status. —C ybianthus vasquezi1 occurs in primary serra
firme lowland forest, with scattered white sand areas. Label data do not al-
low determination of whether this species occurs on the laterite or on the
sands. Because it is known only from two gatherings, its conservation sta-
tus is unknown. However, its importance as a tonic in Mayna Jivaro cul-
ture may indicate it is locally common.

Etymology.—lIt gives me great pleasure to dedicate this species to a great
friend and colleague, Ing. Rodolfo Vasquez Martinez, Assistant Curator and
Director of the Flora of Peru Program of the Missouri Botanical Garden.
Vasquez is an indefatigable collector, a forestry engineer, dendrologist, taxonomist
and author of numerous publications on uses of Peruvian forest products,
economic plants of the Peruvian Amazon, and Florala of the Biological Re-
serves of Iquitos. He is a taxonomic authority on Caraipa and Myristicaceae
of the Amazon Basin.

Local names and uses.—Petu: “sésa,” “kurtip” (Mayna Jivaro). The sap is

extracted and a juice is drunk to “improve hunting ability.”
PARATYPE: PERU. Loreto: Prov. ony oe Hermosa and vicinity, Rio Corrientes, |
m S of junction with Rio Macusari, 03° 15'S, 75° 50' W, 160 m, 3—20 Dec 1985 (pist.
fl), W. Lewis et al. 10306 (BRIT, MO).

Cybianthus vasquezii is most closely related to C. potiaei of the eastern Amazon
Basin (French Guiana and Brazil (Amapa, Bahia), but is easily recognized
by the conspicuously rubiginous punctate-lineate branchlets, larger
pseudoverticillate, oblanceolate leaves, longer petioles and inflorescences,
ovate, apically rounded corolla lobes that are abruptly constricted basally,
and coarsely crenate along the margins.

20. aes cenepensis Pipoly, os nov. (Fig. 20). Typr: PERU. Amazonas:
o Cenepa, ieee Huampami, ca. 5 km E of Chavez Valdivia, 04° 30'S, 78° 30
- 200-250 m, 12 Aug 1978 (stam. fl), ms Kijikat 205 (HOLOTYPE: MO; isoryPes: F,

MO, NY, US, . M).

Ol folia charracea oh] ] ]

ad bases cuneata, petiolos canalicuatos,
inflorescentias anguste pina ad C. buchtieni valde affinis sed ab ea ramulis
alatis (nec laevibus), petiolis 2—2.5 (non 1.5—1.8) cm longis, lobis calycinis late ovatis (nec

Pipoty, Cybianthus in Ecuador and Peru 107

Fic. 20. Cybianthus cenepensis Pipoly. A. Habit, showing winged branchlets. B. Staminate
inflorescence branch, showing secondary inflorescence branch bracts. C. Staminate flower,
showing stamens subequal to corolla lobes, conspicuous square staminal tube, and coarsely
crenate corolla lobes. D. Pistillate flower, showing ellipsoid saul E. Pistillate inflorescence
branch, ene dense spike appearing glomerulate. D. Branchlet apex, showing puberu-

lent vestiture. A drawn from holotype. B—D, drawn from Ancuash 522. E-F, drawn from
Kujikat 306. ‘Figure drawn by Linda Ellis.

108 Sipa 18(1)

lineari-lanceolatis), grosse crenatis (nec integerrimis), lobis corollinis grosse crenatis (nec
enteris) denique antheris ad apices acutis (nec rotundatis) ad bases cordatis (nec obtusis)
perfacile distinguitur.

Treelet to 2 m tall. Branchlets prominently longitudinally ridged, the ridges
forming small, rounded wings, (2.5—3—)5—6 mm diam., sparsely rufous
puberulent, glabrescent. Leaves alternate; blades chartaceous, oblanceolate,
(13.7-)18.5—23(-26.2) cm long, (4-)6-10 cm wide, apically acuminate,
basally cuneate, glabrous above, sparsely rufous puberulent below, glabres-
Cent, an pellucid punctate, the margin entire, flat; petioles

canaliculate, 2—2.5 cm long, sparsely rufous puberulent, glabrescent. Staminate
inflorescence a iS bipinnate panicle, 14-18 cm long, 1—1.7 cm wide, the rachis
densely rufous papillate; secondary inflorescence bracts chartaceous, linear-
lanceolate, 44.5 mm long, 1—1.2 mm wide, apically subulate, densely rufous
tomentellous, the margin irregular, entire; inflorescence branches 3-8 mm
long, the flowers densely subspicate, appearing glomerulate; floral bracts
chartaceous, linear, 1-1.2 mm long, 0.1—0.2 mm wide, apically subulate,
densely glandular-papillate, the margin crenulate, glabrous; pedicels cylindrical,
0.2—0.3 mm long, densely glandular-papillate. Staminate flowers 4-merous,
chartaceous, brownish-purple, 1.6-1.8 mm long; calyx cotyliform, 1-1.2
mm long, the tube 0.3—0.4 mm long, the lobes broadly ovate, 0.7—0.8 mm
long, 0.6-0.7 mm wide, apically acute, prominently brown punctate, the
margin hyaline, coarsely crenate, glabrous; corolla subrotate, 1.2—1.3 mm
long, the tube ca. 0.1 mm long, densely glandular-granulose within, the
lobes widely ovate, 1|—1.2 mm long, 0.8-1 mm wide, apically subacuminate,
glabrous and inconspicuously orange punctate without, glabrous within except
under the filaments, the margin hyaline, coarsely crenate, glabrous; stamens
subequalling the corolla lobes, 1—1.1 mm long, the tube conspicuous, carnose,
square, 0.1—0.2 mm long, densely glandular-granulose, the filaments 0.6—
0.7 mm long, the anthers ovate, 0.3—0.4 mm long, 0.4—0.5 mm wide, apically
acute, basally cordate, the connective inconspicuously orange punctate; pistillode
obturbinate, 0.1—0.2 mm long, 0.2—0.3 mm diam., hollow, glabrous. Pis-
tillate inflorescence as in staminate but 12—14 cm long, 0.8—1.0 cm wide; secondary
inflorescence bracts 3.84.1 mm long, 0.8—1 mm wide; inflorescence branches
3.5-6 mm long; floral bracts 1—-1.2 mm long, 0.1—0.2 mm wide; pedicels
0.2—0.3 mm long. Pest///ate flowers as in staminate but white, 1.1—-1.4 mm
long; calyx 0.6—0.8 mm long, the tube 0.1—0.2 mm long, the lobes 0.5—
0.6 mm long, 0.3-0.4 mm wide; corolla 0.7—0.8 mm long, the tube ca.
0.1 mm long, the lobes 0.6-0.7 mm long, 0.5—0.6 mm wide; staminodes
much sorter than the corolla lobes, 0.3-0.4 mm long, the tube conspicu-
ous, carnose, circular, ca. 0.1 mm long, the filaments ca. 0.1 mm long, the
anthers 0.2—0.3 mm long, 0.1—0.2 mm wide; pistil ellipsoid, 0.5—0.6 mm
long, 0.2-0.3 mm diam., the style 4-lobed, the lobes curved distally, pla-

—

Pipoty, Cybianthus in Ecuador and Peru 109

centa subobose, bearing 4 partially immersed ovules. Fruit globose, 6—7
mm long and in diam., prominently red punctate.

Distribution.—Known only from the Rio Cenepa Drainage Basin, Amazonas,
Peru, 200-250 m elevation, and from one disjunct population from Napo,
Ecuador, very close to the Colombian border.

Ecology and conservation status. —Cybianthus cenepensis inhabits varzea for-
ests in the Amazon of Ecuador and Peru. Despite lack of collections, it is
known that the Napo and the Cenepa Rivers are shallower than many with
their same volume, thus giving them strong currents. Aside from the need
for much more exploration at the edge of the Amazon Basin in Ecuador and
Peru, perhaps the strong current of these rivers makes the dynamics of their
adjacent varzea forests different from others in Amazonia. At this time, the
conservation status of this species is unknow

Etymology.—The specific epithet refers to ~ region of Peru from which
the type was collected, the Rio Cenepa Drainage Basin of the Alto Amazonas
Province, Amazonas Department, Peru.

Local names and uses. —Peru: “sauka” (Huambisa).

Pararypes. ECUADOR. Napo: Cantén Orellana, Yasunf National Park, Maxus Rd. and
pipeline construction project, km 15, 01° 31'S, 76° 32' W, 250 m, 30 Jun 1994 (bud), N.
Pitman 461 (BRIT, MO, QCNE); Rio Yasuni, peri riodically inundated forest ca. 80 km upeNe:
from Nuevo Rocafuerte, 225 m, 17 Sep 1977 (fr), R. Foster 3708 (F, QCA). PERU. Amazonas
Quebrada Sasa, Rfo Cenepa, 250 m, 2 Jun 1973 (stam. fl), E. Ancuash 522 (MO, USM):
Rio Cenepa, vicinity of Huampami, ca. 5 km E of Chavez Valdivia, ca. 4° 30'S, 78° 30' W,
200-250 m, 12 Aug 1978 (pist. fl), A Kujikat 306 (F, MO, USM); eee Chigki Shiunk,
4° 30'S, 78° 30' W, 11 Aug 1978 (bud), E. Ancuash 1412 (MO, USM).

Cybianthus cenepensis is closely related to C. buchtient Pax of the Mapiri
region in Bolivia. However, Cybianthus cenepensis is easily recognized by its
winged branchlets, petioles 2—2.5 cm long, widely ovate and coarsely crenate
calyx lobes, coarsely crenate corolla lobes, and anthers with acute apices and
cordate bases. The ellipsoid pistil is also unique within the subgenus.

27. Cybianthus nanayensis (J.F. Macbr.) G. Agostini, Acta Biol. Venez.
10:160. 1980. Weigeltia nanayensis J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser.
11:33. 1931. Type: PERU. Loreto: Lower Rio Nanay, 4 Jun 1929 (stam. fl), LZ. Williams
658 (HOLOTYPE: F; fragment, G

Weigeltia silvestris J.F. Macbr., Candollea 6:16. 1934. syn. nov. Cybianthus silvestris J.F.
Macbr.) G. Agostini, Acta. Biol. Venez. 10:163. 1980. Type: PERU. Loreto: Mishuyacu,
near Iquitos, Dec 1929 (stam. fl), G. K/vg 724 (HOLOTYPE: F; isoryPEs: NY, US).

hes dubia J.F. Macbr., Candollea 6:17. 1934. syn. nov. Cy Meas PUG E Macbr.)

tini, Acta Biol. Venez. 10:158. 1980. Type: PERU. Loreto: Mishuyacu, near
a 24—28 Sep 1929 (pist. fl, fr), A. Killip G A. fe ooo {OLOTYPE: F; ISOTYPES:
NY, US) {erroneously cited as G. Klug 29900].

Conomorpha loretensis Lundell, Wrightia 6:113. 1980. syn. nov. Cybranthus loretensis (Lundell)
Pipoly, Brittonia 33:496. 1981. Type: PERU. Loreto: Quistococha, near Iquitos, 18
Nov 1977 (stam. fl), A. Gentry 20763 (HOLOTYPE: LL-TEX; isotypes: F, MO, NY).

110 Sipa 18(1)

Subshrub to shrub to 0.5(-1.5) m tall. Branchlets 0.2—0.5 cm thick, smooth,
rufous-lepidote. Leaves in loose pseudoverticels; blades chartaceous to sub-
coriaceous, elliptic, lanceolate or oblanceolate, (7—)11—18(—25.5) cm long,
(2.9-)3.5-7.5(-10) cm, apically acute to acuminate, basally acute to cu-
neate, midrib depressed above, prominently raised below, the secondary veins
7-16 pairs, glabrous above, sparsely and minutely rufous-lepidote below,
puncticulate, the margin flat, entire; petioles marginate, (0.6-)0.9-1.5 cm
long, glabrous. Inflorescences monomorphic, a simple raceme or rarely with a
second, malformed basal branch, (1.5—)3—9.5(-14.5) cm long, the rachis
densely glandular-papillate; floral bracts membranac eous, linear-lanceolate,
0.2—1.2 mm long, 0.1—0.2 mm wide, the apically long-attenuate, sparsely
glandular-papillate, the margin glandular-ciliate, entire; pedicels cylindri-
cal, (0.7—)1-1.5(—2) mm long, densely glandular-papillate. Staminate flowers:
white to cream, 4-merous, chartaceous; calyx cotyliform, 1-1.3 mm long,
the tube ca. 0.1 mm long, the lobes widely ovate to deltate, 0.9-1.2 mm
long, 0.8-1.2 mm wide, apically acute to acuminate, medially thickened
and prominently black punctate, glabrous, a margin hyaline, irregular,
densely glandular-ciliolate; corolla rotate, 1.9—2.3 mm long, the tube 0.6—
0.8 mm long, the lobes widely ovate to ovate, 1.3—1.6 mm long, 1.2—1.4
mm wide, apically acute to rounded, subentire or ee notched, medi-
ally thickened and prominently black punctate, glabrous without, glandu-
lar-granulose within at junction of tube and lobe, the margin hyaline, ir-
regular, glabrous; stamens 2.2—2.5 mm long, exserted or rarely subequal to
corolla, the staminal tube carnose, conspicuous, ().5—0.7 mm long, subtruncate

between the filaments, the apically free portions of the filaments terete, (1.2—)
1.4-2 mm long, the anthers widely ovate, 0.32—0.36 mm long, apically
rounded to obtuse, basally cordulate, the connective prominently punctate
dorsally; pistillode vestigial, obclavate or tubiform, 0.7—0.8 mm long, sparsely
translucent glandular-lepidote, hollow. Pisti//ate flowers as in staminate but
beige to brown; calyx 2—2.1 mm long, the tube 0.1—0.2 mm long, the lobes
1.9-2 mm long, 1.9—2 mm wide; corolla cotyliform, 1.9—2 mm long, the
tube 0.5—0.6 mm long, the lobes widely ovate, 1.2—1.4 mm long, 1.3—1.4
mm wide, apically acute to acuminate; staminodes resembling stamens, 1.2—
1.3 mm long, the staminodial tube 0.5—0.6 mm long, the filaments 0.4—
0.5 mm long, the antherodes subquadrate, ca. 0.2 mm long and wide, apically
obtuse, basally truncate; pistil ellipsoid, 1.4-1.6 mm long, the ovary 1—
1.2 mm long, 1.1—1.2 mm diam., densely translucent glandular-lepidote,
the placenta cupuliform, ovules 2, partially immersed, the style trunctae,
0.3—0.4 mm long, the stigma punctiform. Fruit depressed-globose, 4.5—
5.5 mm long, 5.5—6.5(—7) mm diam., yellow when fresh, prominently black
punctate.

Prpo.y, Cybianthus in Ecuador and Peru 111

Distribution.—Endemic to the tall moist forests on white sands of the
Peruvian Amazon, primarily from the Iquitos area, 1OO—160 m

Ecology and conservation status. —C ybianthus nanayensis is locally common,
and thrives in gaps left by large treefalls in overmature forests, and along
the margins of forest margins and paths, where it occurs in a rather dense
herbaceous layer. However, it does not tolerate compacted soils. With de-
creasing quantitites of habitat owing to logging pressures, the species should
be considered threatened. As a gap species growing on nutrient deficient
soils, and with very attractive fruits, Cybianthus nanayensis shows great promise
as a potentially marketable horticultural plant.

Etymology.—The epithet takes its name from the river basin where it occurs,
the Nanay River.

Representative specimens examined. PERU. Loreto: Prov. Loreto, Nauta, 04°32'S, 73°35!
W, 160 m, 2 Jun 1984 (stam. fl), R. Vasquez & N. Jaramillo 5075 (AMAZ, MO, NY), (fr),
R. Vasquez & N. Jaramillo 5086 (AMAZ, MO); Prov. Maynas, Allpahuayo, IIAP Experi-

mental Station, 04° 10'S, 73° 30' W, 120 m, 20 Sep 1990 (fr), J. Pipoly et al. 12263 (AMAZ,
MO, USM), 15 Aug 1990 (fr), R. Vasquez & N. Jaramillo 14204 (AMAZ, MO, US, USM),
10 Oct 1990 (stam. fl), R. Vasquez & N. Jaramillo 14465 (AMAZ, MO); Laguna Quistococha,
15 km SW of Iquitos, 8 Jul 1977 (stam. fl),.J. Solomon 3466 (LL-TEX, MO); Mishiana, 30
km SW of Iquitos, Callicebus Biological Reserve, Rio Nanay, 4 km S of Mishana, 19 Aug

1978 (stam. fl), R. Foster 4243 (AMAZ, F), 16 Aug 1980 (stam. fl), 4327 (F-2 sheets); ee
Mishana, between Rio Nanay and Rio Itaya, 130 m, 29 Nov 1977 (fr), A. Gentry et al.
21033 (F, MO); Between Iquitos and Sta. Maria de Nanay, 180 m, 31 May 1978 ee fl),
A. Gentry et al. 22367 (AMAZ, F, MO); 03° 50'S, 73° 30! W, 25 Feb 1981 (fr), A. Gentry
et el. 31479 (AMAZ, MO), 31 Dec. 1982 (mixed- stam. fl, fr), A. Gentry & L. Emmons 38776
(MO-2 sheets), 5 Aug 1990 (fr), R. Vasquez et al. 14161 (AMAZ, MO, USM); Rio Nanay,
May—Jun 1929 (stam. fl), Ld. Williams 657 (F); Mishuyacu, near Iquitos, 100 m, Oct-Nov
1929 (stam. fl), G. Klug 304 (F), May—Jun 1930 (stam. fl), G. Klug 1355 (F); 6 Oct. 1982
(stam. bud), R. Vasquez & N. Jaramillo a MO) 20 Jan 1985 (stam. fl), 25 (MO);
Puerto. Almendras, 03° 48'S, 73° 25' W, 122 m, 4 Jan. 1986 (pist. fl), R. Vasquez G N.
Jaramillo 7070 (AMAZ, MO), 30 May oe bud), 7593 (MO), 20 Oct. ae (fr), 8073
, MO); Roca Fuerte (Momé6n), Oct-Nov 1984 (stam. fl), R. Vasquez G N. Jaramillo
5241. Madre de Dios: Prov. Tambopata, Tambopata Reserve, Rfo Tambopata at mouth of
Rio D’Orbigny, 250 m, 6 Mar 1981 (fr), A. Gentry G K. Young 32025 (AMAZ, MO, NY).

Agostini (1980) had not seen the type specimen of Weigeltia silvestris and
included it in Cybzanthus subgenus Comomyrsine, probably due to matching
witha herbarium misidentification of a specimen of Cybzanthus kayapit (Lundell)
Pipoly as Weigeltia silvestris in NY. I previously recognized Cybianthus dubius
and C. /oretensis as distinct taxa (Pipoly 1981) on the basis of quantiative
characters, and had not seen the type of C. sz/vestris. The exserted stamens
and obclavate or tubiform pistillode of the staminate flowers, and truncate
style with punctiform stigma in the pistillate flowers are unique features
within the subgenus.

112 Sipa 18(1)

Populations corresponding to the type of We/geltia silvestris have slightly
larger leaves, but are otherwise indistinguishable from the type of We/ge/tia
nanayensis. The type of Conomorpha dubia is notable only for differences at-
tributable to the fact that it is a pistillate fruiting specimen. Fieldwork in
Peru has shown that populations corresponding to the type of Conomorpha
/oretensis grow in full sun, and consequently have narrower leaves with longer
petioles than those of the type of Wezgeltia nanayensis.

VII. arcuate, ene Grammadenia (Benth.) Pipoly, Mem. New York
3:47. 1987. Grammadenia Benth., Pl. Hartw. 218. 1846. Type Species:
a narginata Benth. = Cybranthus marginatus (Benth.) Pipoly.

Erect or pendent, evergreen, terrestrial, epiphytic or epipetric dioecious,
bisexual, dioecious or monoecious shribs or small trees. Roots diageotropic.
Bark mostly smooth, gray or sometimes brown, cracking transversely on
older parts. Trwnks distinguishable, normally more or less terete, leptocaulous
or rarely pachycaulous basally, growth dynamics corresponding to Rauh’s
Model (Hallé et al. 1978). Branchlets thin to moderately thick, terete or ridged,
smooth to verruculose to verrucose, glabrous or rarely glandular-papillate
apically, glandular-papillate at first in the leaf axils then glabrescent, with-
out lenticels. Cataphylls and pseudocataphylls absent. Leaves alternate,
supervolute, exstipulate, sessile, acrodromous, apically obtuse-mucronate,
basally auriculate, subamplexicaul, minutely glandular-papillate along midrib
above, glabrescent, glabrous below, hydropotes few or absent above, nu-
merous below, the margins hyaline, membranaceous, prominently punc-
tate and punctate-lineate, entire or r minutely crenulate to denticulate; peti-
oles absent. ne phic, simple, axillary raceme, at times reduced
to appear dichasial; i bract broadly ovate to deltate, acute, prominently
punctate, the margins entire to erose, early caducous; rachis terete, straight
or flexuous, minutely glandular-papillate, glabrescent; floral bracts solitary,
at pedicel base, ovate to lanceolate, persistent; pedicels cylindrical to clav-
ate, minutely glandular-papillate, glabrescent, accrescent in fruit. Flowers
unisexual or bisexual, (4—)S—6(—7)-merous; perianth lobes imbricate or
quincuncial or rarely and aberrantly dextrorsely contorted, promiently, con-
spicuously or inconspicuously punctate and punctate-lineate; calyx cotyliform,
the lobes erect, at times reflexed in fruit, the margins irregular or regular,
entire or minutely crenulate, glabrous or glandular-ciliate; corolla rotate,
bearing a ring of glandular granules at the junction of tube and lobe, the
lobes glabrous without, glabrous or rarely glandular-granulose and smooth
or rugose within, the margins regular or irregular, entire or pUnUt EN crenulate;
stamens and staminodes similar, connate to form a conspicuous,
or carnose tube adnate to the corolla tube, spate or with minute lobes
alternating with the anthers, the anthers and antherodes basifixed, sessile

Prpoty, Cybianthus in Ecuador and Peru PS
or on minute apically free filaments, quadrate or ovate, apically emarginate

or rounded, basally truncate, dehiscent by apical birimose pores, usually

dorsally and rarely ventrally punctate; pistil and pistillode similar, obnapiform,

ellipsoid or umbonate, the ovary terete, lobed or costate, sparsely to densely

translucent-lepidote, the placenta umbonate, (1—)2—3(—4) ovulate, the ovules

half-immersed in the basal placenta, uniseriate or biseriate, the style short,

truncate, the stigma punctiform. Frit drupaceous, depressed globose, obovoid,

or ellipsoid, the exocarp sometimes fleshy, the mesocarp and endocarp stony,
prominently punctate-lineate, white, lavender or purple-black at maturity,
one seeded, the testa corrugate, the embryo cylindrical, transverse.

A subgenus of 7 species, in the Lesser Antilles, in Mesoamerica from northern
Costa Rica through Panama, in the Andes from Venezuela to southern Peru,
east through the Guayana Highland and to the Serrania de Turumiquire,
Anzoategui, Venezuela. In Ecuador, two species occur. In Peru, 3 species
have been recorded, of which one, Cybianthus lineatus (Benth.) Pipoly, for-
merly thought to be a Guayana Highland endemic (Pipoly 1987) is reported
for the first time here.

KEY TO THE SPECIES OF CYBIANTHUS SUBGENUS GRAMMADENIA
1. Branchlets verruculose- to verrucose-papillate or red glandular-papillate; leaves
coriaceous, symmetr rical; inflorescence erect, flexuous; perianth coriaceous;

epipetric or terrestrial shrubs or trees of lagunas in paramos and elfin forests

rin “jalca” or “pajonal” alpine savannas.

2. Epipetric se or tree to 6 m tall; crunk ler
pillate prominenily ridged, rugose when dried; leaves oblong,

i] } hl |
VELILUCULOSE

to verrucose-p
elliptic or tnceolate: (l. 1—)1.4—2(—2.5)cm wide; fruit ovoid, then ellip-
soid at maturity; plants of paramos or elfin forests. .........0cce 28. C. marginatus

. Terrestrial shrub to 1.5 m tall; trunk pacnyceuinns, eres oy: glan-
dular-papillate, terete, smooth when dried; | lat 4—)0.6—
1(-1.2) cm wide; fruit obovoid throughout development; eee of j es
or pajonal alpine savannas.
. Branchlets smooth, glabrous; leaves chartaceous, asymmetrical; —
lax, straight; perianth chartaceous; epiphytic shrub or tree in cloud forests

below subpdramos or facultative epiphytes in montane and elfin forests.
30. C. magnus

9. C. lineatus

me

3. Leaves not bearing hydropoten above, conspicuously black punctate and
nctate-lineate below; er ene rachis pac pune: lineate; Be eri-

pu
anth whitish-green, prominently bl
ovate, rounded apically; branchlets fe 7mm diam: fruit purple- black at
30a. C. meus ae magnus
ll

urity.
| . i ] kK ] 4 1]
pe

to orange punctate: inflorescence rachis orange punts: -lineate: an
maroon to purple, rarely white (then pistillate), prominently orange-punctate;
anthers quadrate, emarginate apically; branchlets 2.5—3.5(—4.0) mm diam.,
fruit white, chen lavender at maturity. ....... 30b. C. magnus subsp. asymmetricus

Sipa 18(1)

ee
om! ~
Aste Ae
&

ae
=

iN

' iy
f's
-
Al

;

Cybianthus marginatus

iene

!
90 80

Fic. 21. Pictorialized distribution of C. marginatus. A-K, Variation in leaf shape, marginal
venation and es note prominent apical mucron, sessile leaf base typical of subge-
K, drawn from: A. Cuatrecasas oF B. Pipoly 6954, C. Pipoly 6539,
D. Liesner 8038, E. nee 9032, F. Steyermark 100867, G. Mason 13730, H. Pipoly 6975 ,
I. Pearce 250, J. Lehmann 599, K. Luteyn 10175. Bipare ai Pipoly, 1987, drawn by Bobbi
Angell.

28. ea marginatus ae ) Pipoly, (Fig. 1LA,B, 7C,F, 21). Mem.
ork Bot. Gard. 43: 987. Grammadenia Eman ater Benth., Pl. Hartw.

at ne Type: COLOM a nen Near Pit 3,636 m, 1843 (bisex. fl), C.
Hartweg 1200 (HOLOTYPE: K; isorypEs: BM, E, G- are G-DEL, LD, OXF, P, W-2

Grammadenia Hae oie Mez in Engl., Pfanzenr. IV. 236(Heft 9): 231. 1902. Type:
ones M AMA: Altos de Otesas, 3,300 m, 11 Jan 1883, (bisex. fl), F Lehmann
S).

2399 anaes by Pipoly 1987: G; IsoLecrorypEs: LE, U

Pipoty, Cybianthus in Ecuador and Peru 115

ee alpina Mez in Engl., Pflanzenr. IV. 236(Heft 9):231. 1902. Type: VEN-
EZU A. Andes of Trujillo and Mérida, 1,212—4,390 m, 1842 (bisex. fl), J. Linden
447 (LECTOTYPE by geeRs 1987: P; 1sotecrotypes: BM-2 sheets, BR, G, G-DEL, G-
BOISS; Ky OXE S, V

alee pastensis ae in Engl., Pflanzenr. IV. 236(Heft 9):232. 1902. Type: CO-
LOMBIA. NariNo: W cordillera of Pasto, 3,000—3,200 m, 20 Feb 1881 (bisex. fl), F.
Lehmann 599 (HOLOTYPE: G; ISOTYPES: BM, LE).

Grammadenia nitida Mez in Engl., Pflanzenr. IV. 236(Heft 9):232. 1902. Type: PERU.

<NUCO: Pozuzo, 2,131—2,727 m, 1863 (bisex. fl, fr), R. Pearce 250 (HOLOTYPE: K).

Grammadenia weberbaueri Mez, Repert. Spec. Nov. Regni Veg. 16:418. 1920. Type: PERU.
Cajamarca: Jaén, cordillera E of Huancabamba, E slopes, 2,400-—2,500 m, Apr 1912
(bisex. fl), A. Weberbaner 6121 (Lecrotyre by Pipoly 1987: GH; IsoLectortyPe: F).

Grammmadenia Wea Pittier, J. Wash. Acad. Sci. 21:140. 1931. Type: VENEZUELA.
Meéripa: Tabay, 2,500—3,000 m, 18 Sep 1930 (bisex. fl), W. Gebriger 471 (HOLOTYPE:
VEG ISOTYPES: - F, G, NY, PH).

Gran nia andicola Cuatrec. Seg eee Colomb. Ci. Exact. 8(41):321. 1951.
COLOMBIA. VALLE: cortillets Occidental, Los Farallones, NW slope, Onebiad
Nieves, below El Diamante, 2,900 m, 30 Jul 1946 (bisex. fl, fr), J. Cuatrecasas ae
(LECTOTYPE by Pipoly 1987: F; tsotecrorypes: F, COL 2-sheets, U, US).

Epiphytic shrub or small tree to 6 m, the trunk leptocaulous. Branchlets
prominently ridged, 3-4(-5) mm diam., verruculose- to verrucose-papil-
late, rugose when dried, glabrous. Leaves coriaceous, symmetrical, oblong,
elliptic or lanceolate, (3—)3.5—6(-6.5) cm long, (1.1—)1.4—2(-2.5) cm wide,
apically acute to obtuse, mucronulate, basally acute, auriculate, nitid above,
pallid below, prominently punctate and punctate-lineate, the margin en-
tire, revolute. Inflorescence erect, the rachis flexuous, slender, (1—)1.3—2(—3.2)
cm long, densely black punctate-lineate; floral bracts widely ovate to deltate,
(0.8—)1.2—1.5(-1.8) mm long, 0.8—1.4(-1.8) mm wide, apically acute, the
margin entire; pedicels (1—)1.5—2 mm long. Flowers coriaceous, 5(—7)-merous;
calyx 1.1—2.1 mm long, the tube ca. 0.1 mm long, the lobes deltate, (1—)
1.5-2 mm long, (1.2—)1.3—1.5(-2) mm long, apically acute, prominently
and densely punctate and punctate-lineate, the margins minutely crenu-
late, glabrous to minutely ciliolate at first apically; corolla (2—)2.2-3 mm
long, the lobes widely ovate, (1—)1.2—2 mm long, (1.2—)1.5-2(-2.5) mm
wide, obtuse to rounded, glabrous without, glandular-granulose within basally,
densely and prominently punctate medially, the margin entire; staminal tube
carnose, 0.6—0.8(—1) mm long, the anthers sessile, alternate with fleshy lobes
to 0.2 mm long, quadrate, 0.4—0.7 mm long and wide, apically rounded,
basally truncate, the connective prominently punctate dorsally; pistil obnapiform,
(0.9-)1.2-1.5 mm long and (0.8—)1.2—1.6 mm diam., the ovary (0.4—)0.6—
1 mm long, glabrous to sparingly translucen glandular-lepidote, ovules 2(—
4), uni- or biseriate, the style 0.3-0.5 mm long, epunctate. Fruzt somewhat
ovoid, then ellipsoid at maturity, 4.5-5(—6) mm long, (2.5—)3—4 mm diam.,
white, then purple-black at maturity, the punctations red-black, prominent.

Distribution. —Cybianthus marginatus is the most common species of the

116 Sipa 18(1)

subgenus, occurring throughout the Andes from Venezuela to Peru, 2,000—
3,400 m.

Ecology and conservation status. —This species occurs in large populations
on rocks above lagunas in paéramos in northern Ecuador and along water-
courses in paramoid elfin forests, and “ceja” formations in the remainder of
Ecuador and Peru. As long as there are remnant péramo formations, or elfin
forests, there will be populations of Cybianthus marginatus. However, in some
instances, soil compaction due to overgrazing by sheep can render the soil
uninhabitable for this species.

Etymology.—tThe specific epithet refers to the scarious leaf margin, made
conspicous by the coriaceous texture. In addition, this aspect is made more
conspicuous by the prominent submarginal vein of many populations.

Representative specimens examined. ECUADOR. Azuay: “Oriente” border, Paramo del
Castillo, crest of E cordillera on trail between Sevilla de Oro and Méndez, 2,727—3,333 m,
18 Aug 1945 (fl, fr), W. Camp E-4809 (NY, VEN); Rio Collay, Sa ea s . . Pan,
2,650—3,290 m, 6 Jul ene }: Stojnik 53854 ON). carchi: Peak of C
00° 51' 12" N, 78° 08' 21" W, 3070 m, 24 Jul 1994 (fr), B. Boyle et al. 3373 (BRIT. ‘MO.
QCNE); Cantén eked Loma El Coraz6n, Bretafia, SE of Mariscal Sucre, Rfo Minas,
00° 35' N, 77° 42' W, 3,150 m, 22—23 Dec 1992 (Al, fr), W. Palacios & G. Tipaz 10569
(BRIT, MO, QCNE). Imbabura: Cordillera Oriental, Camp Arelan, E of Volcin Cayambe,
2,803 m, 21 Jul 1944 (A, fr), W. Drew E-351 (MSC); Ridge just § of Rio Clavadero, along
trail co Rio San Pedro, E of Cayambe, 2,893 m, 27 Jul 1944 (A, fr), 1. Wiggins ae (DS,
US). Loja: Saraguro-Loja, Km 12.4, turnoff toward Fierro Urco, Km 2.5—2,7, 03° 41! 05"
S, 79° 17! 20" W, 3,150—3,300 m, 7 Dec 1994 (fl), P Jorgensen et al. 1278 (BRIT, LOJA,
MO, QCA, QCNE); Paramos de Saraguro, 10 km S of Saraguro, 3,050 m, 2 Jan 1979 (fl),
J. Luteyn et al. 6647 (NY, QCA). Zamora-Chinchipe: border, crest of Cordillera Oriental,
2,840 m, 28 Jan 1984 (fl), J. Luteyn & E. Cotton 11295 (NY, QCA); W slopes of ee
del Condor and NW slopes of Nudo de Sabanillas, around Tambo Cachiyacu, ca
of Yangana, 2,000—3,000 m, 19 Oct 1943 (A, fr), Ji SLCjermate 54800 (NY, U). S of
Play6n de San Francisco, slopes of Cerro Mirador, 3,300—3,600 m, 29 Dec 1980 (fl, fr), L.
Holm-Nielsen et al. 29949 (AAU), J. iia et al. 3 929 (AAU, QCA). PERU. Amazonas:
Prov. Luyas, Dtto. Camporredondo, Anexo Tullanaya, Cerro Wicsocunga, 06° 05' 35" S,
78° 19' 56" W, 3,075 m, 7 Dec 1996 (pist. fl), J. & L. Campos 3121 (BRIT, MO, USM).
Cajamarca: Jaén, SW of Querocotillo, 3,150 m, Aug 1915 (bisex. fl, fr), A. pe
7168 (F, G, GH). Cusco: La Convencién, 2,800 m, 9 Jul 1968 (bisex. fl, fr), ’ Dudley
10910 (NA), 10 Jul 1968 (bisex. fl, fr), [7 Dudley 10922 (F, NA), T. Dudley ie (F,
NA). Huanuco: Prov. Hudnuco, 45 km on rd. from Hudnuco to Tingo Maria, trail on $
side of Carpish Tunnel, 09° 42'S, 76° 05' W, 2,400 m, 3 Mar 1985 (bisex. fl), C. Todzia G
B. Stein 2740 (F, TEX, USM).

jon

As was stated previously (Pipoly 1987), Cybianthus marginatus is most
closely related to C. /ineatus (Benth.) Pipoly, previously known only from
the contiguous Guayana Floristic Province, because of its (synapomorphic)
flexous inflorescences and biseriate ovules. However, Cybianthus marginatus
is easily distinguished from C. /ineatus by its glabrous, ridged, verrucose-
papillace branchlets, oblong, elliptic or ovate leaves, obnapiform pistil and

PipoLy, Cybianthus in Ecuador and Peru 117

ellipsoid fruits. The verrucose-papillate branchlets, leaf and fruit shape are
unique (autapomorphic) features within the subgenus.

Cybianthus marginatus is the most variable Species of the subgenus, con-
taining one-third of the names attributed toG synonyms
Variation in leaf size and punctation and quantitative floral variation have
led to much overdescription. A full discussion of synonymy and variation
was provided by Pipoly (1987). Cybianthus marginatus is most closely re-
lated to C. /ineatus by virtue of its flexuous inflorescences and biseriate ovules.
However, Cybianthus marginatus is easily distinguished from C. /zneatus by
the glabrous, ridged, verruose-papillate branchlets, oblong, elliptic or ovate
leaves, obnapiform pistil and ellipsoid fruits.

on

29. Cybianthus lineatus (Benth.) Pipoly (Fig. 7E), Mem. New York Bot.
Gard. 43:64. 1987. Grammadenia lineata Bentham, Pl. Hartw. 218. 1846. Type:
Ae [“GUYANA’}. BoLivar: Savannas near Rone 1843 (bisex. fl, fr), R.
Sc ; 47/992 (HOLOTYPE: K; 1isoryPEs: B, BM, G-DC, G-DEL, P, U, W-2 sheets).

Poe shrub to 1.5 m tall, the trunk pachycaulous. Branchlets terete,
(2.5—)3—4(—6) mm diam., densely red glandular-papillate apically. Leaves
coriaceous, symmetrical, oblanceolate, (1.2—)1.6—3(—3.9) cm long, (0.4—)0.6-
1(—1.2) cm wide, apically acute, basally subauriculate, the margin flat. Inflorescence
erect, flexuous, at times reduced to a simple dichasium, 0.6—1.0(—2.5) cm
long, sparingly glandular-papillate; floral bracts widely ovate, 0.8—1.1 mm
long, 1.1—1.4 mm wide, apically acute, the margin entire, glabrous. F/ow-
ers chartaceous, 5—6(—7)-merous; calyx 1.1—-1.6 mm long, the tube ca. 0.1
mm long, the lobes wide-triangular to deltate, (0.8—)1—1.5 mm long, (0.7-)
1—1.2(—1.4) mm wide, apically acute, conspicuously black punctate and punctate-
lineate, the margins entire, glandular-ciliolate; corolla (1.8—)2—2.5(—3) mm
long, the lobes widely ovate, (1—)1.3—1.6(—2) mm long, (1—)1.3—1.7(-2)
mm wide, obtuse to emarginate, inconspicuously punctate medially, the
margins irregular, entire; staminal tube carnose, conspicuous, (0.6—)0.8—1
mm long, the anthers sessile, alternate with prominent fleshy lobes, quad-
rate, 0.4—0.6 mm long and wide, apically rounded, prominently black punctate
dorsally; pistil ellipsoid, (0.9—)1—1.2(-1.4) mm long, (0.8—)1—1.3 mm diam.,
the ovary 0.8—1 mm long, glabrous to translucent glandular-lepidote apically,
ovules 2—3, when more than 2, biseriate, the style (O.1—)0.2(—0.3) mm long,
glabrous. Fruit obovoid, (3—)4—5 mm long, (2—)2.5—3 mm diam., purple,
than black at maturity, the punctations green, prominent.

Distribution.—Formerly thought to be endemic to the Guayana High-
land, in open savannas on tepuf summits throughout Pantepui (Mayr & Phelps
1967), at 1,400—2,850 m elevation, but now known elsewhere only from
the collection cited below.

Ecology and conservation status. —C ybianthus lineatus grows in fully exposed,

118 Sipa 18(1)

dry montane “shrub savannas” (Huber 1995) in shallow sand over sand-
stone throughout Panteput. Its occurrence in Pajonal (“jalca”) vegetation
in Peru gives reason to expect it in other places, especially in the Cordillera
del Condor along the Ecuadorean/Peruvian border. Unfortunately, recent
civil unrest has prohibited collection in that area. In the majority of the
range for the species, Cybranthus lineatus is not threatened.

Etymology.—The specific epithet refers to the numerous and prominent
punctate-lineations of the abaxial leaf surface.

Specimen examined. PERU. Pasco: Oxapampa Prov., Cerro Pajonal, 29 km from Oxapampa,
2,680 m, D. Smith & Foster 2509 (FE MO, USM).

Cybianthus lineatus is most closely related to C. marginatus (Benth.) Pipoly,
by virtue of its flexuous inflorescence and biseriate ovules (Pipoly 1987).
However, Cybianthus lineatus may be easily recognized by its pachycaulous
crunk, densely red glandular-papillate branchlet apices, ellipsoid pistil and
obovoid fruits. It is the only species in the subgenus with parenchyma in-
stead of aerenchyma in the cortex, the only one with bifacial palisade layers
in the leaf, and the only one with a pachycaulous trunk. None of these
morphological peculiarities are unexpected given its drier, wind-swept habitat.

30. Cybianthus magnus (Mez) Pipoly, Mem. New York Bot. Gard. 43:55. 1987.

Facultative epiphytic shrub or tree to 7 m tall. Branchlets terete, smooth,

glabrous. Leaves chartaceous, asymmetrical, narrowly oblanceolate, oblan-
ceolate or narrowly obovate, (4.5—)5.2-15 cm long, (1.0—)2.1—5.2 cm wide,
apically acute to abruptly acuminate, tapering abruptly or gradually to base,
bearing hydropotes above or not, conspicuously black punctate and punc-
tate-lineate or inconspicuously pellucid to orange punctate below, the margins
entire, flat, or subrevolute. Inflorescence lax, straight, (1.5—)2—8(-11.5) cm
long, slender, densely glandular-granulose and papillate, prominently black
punctate-lineate or conspicuously orange to brown punctate-lineate; floral
bracts ovate, widely ovate or deltate, (0.7—)1.1—2.2 mm long, (0.6—-)1.3—2
mm wide, apically acute to acuminate, prominently black or orange punc-

tate and punctate-lineate, the margins erose and glandular-ciliate; pedicels
1.0—2.2(—-5.5) mm long in flower, the smaller ones accrescent to 4(—6) mm
long in fruit. F/owers chartaceous, 5(—G)-merous, whitish-green or pink to
maroon; calyx lobes widely ovate to delatate, (0.8—)1—-1.5(—2) mm long,
(0.8—)1—2.1 mm wide, apically acute to acuminate, prominently black punctate
and punctate-lineate or orange to brown punctate, the margins erose to fimbriate
and densely glandular-ciliate; corolla (1.7—)2—2.6(—3) mm long, the stami-
nate and bisexual maroon, the pistillate white, the lobes widely ovate, 1.1—
1.6(—2.2) mm long, (0.9—)1.1—2.6 mm wide, apically obtuse to emargin-
ate, rugose medially within, densely and prominently black punctate and

Pipoty, Cybianthus in Ecuador and Peru 119

punctate-lineate or orange to brown punctate, the margins irregular, en-
tire; staminal tube membranaceous, (0.4—)0.7—1(—1.2) mm long, lobate,
the lobes 0.1—0.2 mm long, the anthers sessile, alternate with the lobes,
ovate to quadrate, (0.3—)0.4—0.6 mm long, 0.4—0.6(—7) mm wide, apically
rounded or emarginate, the connectives epunctate ventrally, prominently
black or orange punctate dorsally; pistil obnapiform, 1—1.2 mm long, 0.9—
1.5 mm diam., the ovary 0.6-0.9(-1.3) mm long, densely translucent glandular-
lepidote, the ovules 2—4(—S), uniseriate, the style 0.3—0.5(—0.7) mm long,
glabrous. Fruit obovoid, 2.5—3.5 mm long, 2—3 mm diam. when dried, pink,
then purple-black or white, then lavender at maturity, prominently black
punctate-lineate or orange punctate and punctate-lineate when dried.

Distribution.—Cybianthus magnus occurs in the Andes of Venezuela southward
to Peru as an epiphyte in the cloud forest zone below subpaéramo thickets
and as a facultative epiphyte in montane and elfin “ceja” forests, from 1,100—
3,500 m. It is also known from the Serrania de Turumiquire, in the states
of Monagas, Sucre and Anzodtegui, Venezuela, at 2,000—2,400 m.

Etymology.—The specific epithet refers both to the large, branchlets, often
appearing succulent, as well as the large leaf size found in some populations.

s shown by Pipoly (1987) Cybzanthus magnus is most closely related to

C. parasiticus (Sw.) Pipoly from the Lesser Antilles by its chartaceous co-
rolla rugose medially within, asymmetrical leaves, and obovoid fruits, but
is easily separated from it by its erose and fimbriate calyx lobes, lobate staminal
tube and sessile anthers. Cybianthus magnus superficially resembles C. marginatus
(Benth.) Pipoly, but may be easily separated by its subsucculent smooth
stems, and obovoid fruits

Both subspecies are known from Ecuador and Peru, with one region of
apparent sympatry on the eastern slopes of the Cordillera Oriental in Ecua-
dor and adjacent northern Peru. However, they appear to be separated by
habitat. In Ecuador and Peru, subspecies magnus appears to be restricted to
closed cloud forests, while subspecies asymmetricus is found in open mon-
tane forest and elfin (“ceja”) forest. The salient features of each are summa-
rized below.

30a. Cybianthus magnus (Mez) P1 poly subsp. magnus (Fig. 8E). Mem.
New York Bot. Gard. 43:56. 1987 gna Mez in Engl., Pflanzenr.
IV. 236(Heft 9):231. 1902. Type: COLOMBIA. SANTANDER DEL Norte: Ocafia to
Pamplona, 2,000—2,500 m, 4 Mar 1879 (fl), W. Kalbreyer 1087 (HOLOTYPE: K).

Grammadenia oxygyna Cuatrec., Revista Acad. Colomb. Ci. Exact 8:321. 1951. Type:
COLOMBIA. VALLe DEL Cauca: Cordillera Occidental, W slope, bank of Rio Digua,
left side, Piedra de Moler, 900—1,100 m, 20 Aug 1943 (fl, fr), J. Cuatrecasas 14947 (LECTO-
tyPE by Pipoly (1987): F NY Neg 12136; isotecrotypEs:COL-3 sheets, F, U, US).

Facultative epiphytic shrub or tree to 7 m tall, 7—15(—30) cm diam., the

120 Sipa 18(1)

canopy often bowl-shaped. Branchlets (4.0—)5.0-7.0 mm diam. Leaves not
bearing hydropotes above, conspicuously black punctate and punctate-lineate
below. Inflorescence rachis prominently black punctate and punctate-lineate.
Flowers with perianth whitish-green, prominently black punctate and punctate-
lineate; anthers ovate, rounded apically, the connectives prominently black
punctate dorsally. Frit purple-black at maturity, prominently black punc-
tate-lineate when dried.

Distribution. —Subspecies magnus occurs from the Serranfa de Turumiquire
(states of Anzodtegui, Monagas, Sucre), and in the Andes, from Venezuela
southward through Colombia to and Ecuador to Peru.

Ecology and conservation status.—Subspecies magnus occurs as an epiphyte
in closed cloud forests, especially those below subpdéramo thickets. It is a
relatively rare, but widely distributed subspecies, and is increasingly en-
dangered owing to habitat destruction.

Etymology.—The epithet refers to the leaf size and stem succulence, a novelty
within the subgenus.

Specimens examined. ECUADOR. Loja: Between Nudo de Sabanillas and Rfo Cachiyacu
at Tambo aloe 3, O00—3,500 17 Oct 1943 (fr), J. Steyermark 53584 (NY). Zamora-

chipe: Nangaritza Cantén, ae crest of Cordillera del Condor, above Pachicutza, on
disputed Peru- Bie nee 04° 06'S, Hi 35' W, 1,800 m, 5 Dec 1990 (stam. fl), D
Neill & W. Palacios 9518 (MO, QCNE). PERU. Cajamarca: Prov. San Ignacio, path at the
border of “La Unién,” oe m, | Nov (scam. fl), C. Diaz & A. Torres 7805 (B
MO, USM).

30b. Cybianthus magnus (Mez) Pipoly wee Sele ae ee Pip

Fig. 8F), Mem. New Y Ore Bo : ele td. 43:57. 1987.G
Men, Bull. Herb. Boissier sér 2, 5:2 1905. ie PERU. Loreto: Cerro de Ponasa,

300 m, Mar 1903 (fr), E. Ule Fae TOTYPE by Pipoly 1987: HBG; IsoLeCTOTYPES
: fen K, L).

Grammadenta macrocarpa Lundell, Wrightia 5:292. 1976. Type: ECUADOR. Napo: 17
km aque, 70-73 km W of Lago Agrio, 1,130 m, 4 Nov 1974 (bisex. A,

fr), A. Gentry 12419 (HOLOTYPE: LL-TEX; Isorypes: MO, on

Facultative epiphytic shrb to 2 m tall, 7 m diam., the canopy open, conical.
Branchlets 2.5—3.5(—4.0) mm diam. Leaves bearing hydropotes above at least
proximally, inconspicuously pellucid to orange punctate below. Inflorescence
rachis conspicuously orange to brown punctate-lineate. F/owers with calyx
greenish-pink, the staminate and bisexial with corolla maroon, the pistil-
late with corolla white and prominently orange punctate and punctate-lineate
lobes; anthers quadrate, apically emarginate, the connectives prominently
orange punctate dorsally. Frit white, then lavender at maturity, prominently
orange punctate and punctate-lineate when dried.

Distribution.—C ybianthus magnus subsp. asymmetricus occurs as an obligate
epiphyte, growing on detritis in open montane forests and elfin forests, from
the Darién of Panama to Cusco, Peru, from 1,000—2,000(—2,700) m.

PipoLy, Cybianthus in Ecuador and Peru 121

Ecology and conservation status. —Subspecies asymmetricus grows as an obli-
gate epiphyte, growing on rocks with deep organic detritis and large, moss-
covered trees. With increasing levels of disturbance, it is being threatened.

Etymology.—The epithet refers to the asymmetric shape of the leaf blades.

Specimens examined. ECUADOR. Carchi: Cantén Tulcdn, Parroquia Tobar Donoso,
Reserva Indigena Aw4, Centro E] Baboso, 00° 53' N, 78° 25' W, 1,800 m, 17-27 Aug
a (fr), G. ae et al. 1741 (BRIT, MO, QCNE); Parroquia el Chical, Centro San Marcos,

° 06' N, 78° 14' W, 900-1,100 m, 20-30 Apr 1993 (fl), P aes i al. 341 (BRIT,
noe Cerro Cie valley bottom ca. 1.5 km NNE of summit, 00° 51' 52" N, 78°
O8' 10" W, 2,750 m, 25 Jul 1994 (stam. fl), B. Boyleet al. 3450 (BRIT, pee MO). PERU.

mazonas: pe Ba agua, Imaza, Nuevo Samaria (anexo de UVT), 18 Mar 1995 C. ae i
al. 7585 (BRIT, HUT, MO, USM). Cusco: La Convencién, Cordillera Vilcabamba,
way between Camps 2 1/2 and 3, 1,980 m, 1 Jul 1968 (bisex. fl, fr), 7; Dudley 70668 a
NA), 1,800 m, 24 Jul 1968 (bicoe fl, fr), 1. Dudley 11324 (NA). Huanuco: Prov. Pachitea,
region of Pucallpa, W part of Sira Mountains and oe Sere ca 24 km SE to 26 km
ESE of Puerto Inca, next to Campamento Pato R 09° 27'S, 74° 46' W, and along crest
after Campamento Peligrosol 600 m, 14 Apr 1 see Go, B, es oe 14 Sere nee
W, WU). Pasco: Oxapampa, Cordillera Yanachaga, Cerro Pajonal, c , 12 km SE of
Oxapampa, 2,700—2,800 m, 7 Oct 1983 (bisex. fl), R. Foster 9013 ae < aan

Cybianthus magnus subsp. asymmetricus is notable for its stems with angu-
lar collenchyma in the pith, well-developed aerenchyma in the inner cor-
tex, and tangential collenchyma in the outer cortex. It is separated from
subspecies magnus by its inconspicuous or orange punctate leaves, orange
punctate-lineate inflorescence rachis and perianth, quadrate, emarginate anthers
and open montane and elfin forest habitat.

Grammadenia macrocarpa Lundell is notable only for its large, bright or-
ange fruits and smaller flowers. I examined one fruit from each of the isotypes
of G. macrocarpa and found an insect larva in each one, accounting for the
size and peculiar morphology.

VII. lee Mart. een Cybianthus. Cybianthus sect. Secs
_in Mart., Fl. Bras. 10:292. 1856. Cybianthus sect. Cybianthoides Miq. in Mart
A ‘Bras. 10: 200. 1856. me Specins. Cybianthus penduliflorus Matt.

Peckia Vell., Fl. Flum. 1:51. 1825., nom. reg.

Terrestrial dioecious shrubs or small trees. Roots positively geotropic. Bark
mostly smooth, gray or sometimes brown, cracking longitudinally on older
parts. Trunks distinguishable, normally more or less terete, leptocaulous,
growth dynamics corresponding to Rauh’s Model (Hallé et al. 1978). Branchlets
thin to moderately thick, terete, trigonal, or ridged, rufous stellate or den-
droid tomentose or rarely, with rufous, subsessile covering lepidote scales.
lls absent. Leaves alternate, or pseudoverticillate,
supervolute, exstipulate, petiolate; blades epunctate or variously black or
red punctate, the punctations at times prominent. Inflorescence a simple, lat-
eral (axillary) raceme; peduncle 1-5 mm long, the rachis straight, minutely

Cataphylls and DSelL val 4
a / It

rT

22 Sipa 18(1)

rufous glandular-papillate, glandular-granulose or lepidote, glabrescent or
persistent; floral bracts solitary, at pedicel basally, ovate to lanceolate, per-
sistent; pedicels cylindrical to clavate, or obconic, minutely glandular-papillate,
glandular-granulose or lepidote, accrescent in fruit. F/owers unisexual or bisexual,

4-merous; perianth lobes imbricate or rarely valvate, prominently, conspicuous!
> y >

or inconspicuously punctate and/or punctate-lineate; calyx cotyliform, the
lobes erect, at times reflexed in fruit, the margins irregular or regular, en-
tire or minutely crenulate, glandular-ciliate; corolla rotate, to subrotate, the
tube short, glabrous or glandular-granulose, at times papillate, the lobes
glabrous without, glandular-granulose and/or glandular-papillate throughout

—

a-

brous, glandular-granulose or papillate along the margins; stamens and
staminodes similar, developmentally adnate to the corolla tube to form an
inconspicuous tube, the stamens thus appearing epi Paro apical free portion
of the filaments present or absent, when present up to 3 times longer than
the anthers, the anthers basifixed, appearing sessile or on minute apically
free filaments, quadrate or ovate, apically emarginate, rounded, acute, api-
culate or truncate, basally truncate or subcordate, dehiscent by apical pores,
the pores at times confluent, the connective epunctate or conspicuously or
prominently punctate, at times glandular-papillate; pistil and pistillode similar,
conic, ellipsoid, or rarely, obturbinate, the ovary terete, lobed or costate,
sparsely to densely translucent-lepidote, the placenta subglobose, (1—)2—
3(—4) ovulate, the ovules half-immersed in the basal placenta, uniseriate or
biseriate, the style long-attenuate, the stigma punctiform, lobed or capi-
tate-lobed. Frait drupaceous, depressed globose, the exocarp sometimes fleshy,
the mesocarp and endocarp stony, prominently punctate-lineate, white, red
or purple-black at maturity, one seeded, the testa corrugate, the embryo
cylindrical, transverse.

A subgenus of 50 species in tropical South America, with the largest
concentration of species in the Amazon Basin and adjacent Guianas, Planalto
and coastal Brazil. Subgenus Cybianthus is by far the most complicated taxonomic
group within the genus and is known from rather material. Collection
of more material is hampered by the fact that the populations studied here-
tofore have revealed population densities lower than any other subgenus.
So far, 13 species are known from Ecuador and Peru, but with additional
collections, we may expect to find Cybianthus lanceolatus Pax, and/or Cybianthus
psychotriifolius (Rusby) Mez, both from nearby Bolivia, in southern Peru.

within, che margins regular or irregular, entire or minutely crenulate, g

KEY TO SPECIES OF CYBIANTHUS SUBGENUS CYBIANTHUS
1. Branchlets subterete to angulate
2. Inflorescence spicate or subspicate; calyx lobes inconspicuously or promi-
nently orange punctate; petioles pulvinate basally.

Prpoty, Cybianthus in Ecuador and Peru 123

3. Leaf blades i 7 ] 1] 4] | a | | At }

or carnose, rounded or acute sipsiteally:

4. Branchlets 4—6 mm diam.; leaf blades coriaceous, nitid above; peti-
oles 2.2—3 cm long; inflorescence spicate, the pedicels obsolete to
0.4 mm long; calyx lobes chartaceous, deltate, the margin entire,
undulate, glabrous. 31. C. incognitus
. Branchlet ae 9 mm diam.; leaf blades membranaceous, dull above;
se 1.7—2 cm long; inflorescence subspicate, the pedicels 0.6—
0.9 mm long; calyx lobes carnose, ovate, the margin crenulate, Gee ular-

oS

ciliat 2. C. minutiflorus
S-Leat bla des perpuncticulose below; calyx lobes ee
subacuminate apically. 33. C. huampamiensis

2. Inflorescence racemose; calyx lobes densely and prominently black punc-
tate or epunctate; petioles tapered, not pulvinate basally

5. Leaves alternate; calyx lobes acute apically, the margins hyaline; an-

ther apically rounded, obtuse or acute apically, the pores not confluent.

6. Pedicels 1.9-—2.2 mm long; flowers erect; calyx carnose, 0.8—1.1 mm

long, the lobes ovate, abruptly constricted age: densely and promi-

nently black punctate, the margin irregularly serrate, punctate-lineate,

glabrous 34. C. granulosus
Gi, Beliccs2 .5—3.5 mm long; flowers nodding; calyx membranaceous,

1.5-1.9 mm long, the lobes deltate to triangular, not constricted

basally, epunctate, the margin entire, epunctate, minutely glandu-

lar-ciliolate. 35. C. flavovirens
] a | ] ies! a ae | F 1] iB -
oy I ne oa J I 1 q
anthers truncate apically, the pores confluent. ...........:000006+ 36. C. venezuelanus

1. Branchlets terete
7. Branc i) thick, (6- )7-10 mm in pee
8. Leaf b]

the lobes acute or Seana

9. Leaf blades nitid and perpuncticulose above, 12—13.5 cm wide, the
secondary veins prominently raised above and below, basally trun-
cate, auriculate; petioles 0.5—1.4 cm long; pedicels 0.3—0.5 mm long.

37. C. grandezii

9. Leaf blades sordid and epunctate above, 2.1—5 cm wide, the second-

ary veins deeply impressed above, prominently raised below, basally

cuneate; petioles 2.1-5 cm long; pedicels 0.8-1.

38. C. jensonii
8. Leaf blades abruptly acuminate apically; calyx coriaceous, the lobes

rounded. 39. C. fosteri
. Branchlets thin, 1.5—3.5 mm in diameter.
10 f blades apically subacumi l

or attentuate, the margin erose, ee ae
11. Leaf blades chartaceous to coriaceous, somewhat to very nitid
above and below, the midrib raised above, decurrent to base
of petiole; petioles short-pulvinate basally.
12. Leaf blades e mae Ge )5.5-7(—10.8) cm wide, apically
long-acumi , the tertiary veins prominently
raised, anaes penaciepanerate below; petioles
canaliculate; pedicels cylindrical in fruit. ..........0..0.0. 40. C. resinosus

124 Stipa 18(1)

Fic, 22. Cybianthus incognitus Pipoly. A. Habit, showing trigonal branchlet. B. Abaxial leaf
surface, ae een minute scales. C. Portion of infructescence, showing deltate calyx lobes
with entire margins. D. Branchlet apex, showing dendroid and stellate tomentum. E. Por-
tion of staminate spike, showing conspicuous staminal tube, and obcordate anthers with
subapical non-confluent pores. A, B, D, drawn from Gentry et al. 22911. C, drawn from
Barbour 2567. ¥%., drawn from holotype, by Linda Ellis.

Pipoty, Cybianthus in Ecuador and Peru 125

Zs Leaf blades very narrowly bl | oblong, 2-A(-5) cm

wide, apically and basally long-attenuate, the tertiary veins
conspicuous, conspicuously black or red punctate and

punctate lineate below; petioles marginate; pedicels obconic

in fruit 41.C. fuscus
11. Leaf blades chartaceous, dull green above and below, the mid-
of Baer a aos be ] | Hee | a | ] 1]
tapering to base, without pulvinus 42. C. cyclopetalus
10. Leaf blades apically acute; ¢ clipe lobes obtuse, the margin crenu
late, long ciliate. 43. C. penduliflorus

31. Cybianthus incognitus Pipoly, sp. nov. (Fig. 22). Typr: PERU. Amazonas:
Rio Santiago Valley, 03° 50'S, 77° 40' W, Quebrada Caterpiza, 2-3 km from Caterpiza
settlement, primary forest, 200 m, tree 9 m tall, 12 Dec 1979 (stam. fl), S. Tungur
289 (HOLOTYPE: MO; IsotypPes: USM, NY’).

Ob folia coriacea oo, desuper nitida subter pallida anthera filamenta 3-plo breviores

C. priewro valde arcte affinis sed ab ea petiolis canaliculatis (nec marginatis), ramulis adpresse
canes et sll (nec € erecte dendroideo- ) tomentellis, pedicellis 0O-0.4 (non 0.8—
1.4) mm is, lobi inconspicue pellucido- (non manifeste atro-) punctatis, lobisw

oe ovatis (nec pubacoreuiadis) pistillodio globoso (nec conico) denique fructu laevi
statim recognitur.

Tree to 9 m tall, at times flowering precociously (P. Barbour 2405). Branchlets
subterete to trigonal, 4-6 mm diam., appressed rufous dendroid and stel-
late tomentose. Leaves pseudoverticillate; blades coriaceous, oblanceolate,
(16—)20—25(—31) cm long, (5—)6.5—8.5(—10) cm wide, apically acuminate,
basally cuneate, decurrent on the petiole, nitid above, pallid below, the midrib
slightly impressed above, prominently raised below, the secondary veins 9—
12(-15) pairs, slightly raised above, prominently raised below, nitid above,
pallid and minutely rubiginous lepidote below, the pellucid punctations
inconspicuous, the margin entire, irregular, flat; petioles cana eae 2.2-
3.cm long, pulvinate, sparsely pubescent at first, glabrescent. Sz
an erect, dense spike, (6—)8—9.5 cm long, the rachis green, sparsely den-
droid pubescent, glabrescent; floral bracts lanceolate, 0.5—1 mm long, apically
attenuate, sparsely pubescent, early caducous; pedicels obsolete to stoutly
cylindrical, O-0.4 mm long, glabrate. Staminate flowers 4-merous, yellow,
chartaceous; calyx cotyliform, 0.6—-0.9 mm long, the tube 0.2 mm long,
the lobes deltate, 0.4—0.7 mm long and wide, apically acute, inconspicu-
ously pellucid punctate, the margins scarious, entire, epunctate, undulate,
glabrous; corolla subrotate, 1.4—1.6 mm long, the tube 0.2—0.3 mm long,
the lobes ovate, 1.2—-1.4 mm long, 0.4-0.6 mm wide, apically acute, gla-
brous without, densely glandular-granulose within, inconspicuously pel-
lucid punctate, the margin opaque, densely glandular-granulose, entire; stamens
1-1.2 ‘mm long, the tube ca. 0.2 mm long, the filaments terete, 0.5—0.6
mm long, slightly reflexed proximally, the anthers obcordate, 0.3—0.4 mm
long, apically apiculate, basally cordate, dehiscent by small, subapical, ovate,

126 Sipa 18(1)

non-confluent pores, the pores extending less than 1/2 anther length, the
connective epunctate, densely and minutely rubiginous glandular-granulose
dorsally; pistillode broadly subglobose, ca. 0.2 mm long, 0.4 mm diam.,
densely yellow glandular-papillate. Prsti//ate inflorescence as in staminate but
9—12 cm long; floral bracts lanceloate, 0.2-0.5 mm long, glabrate; pedicels
obsolete to 0.4 mm long, glabrate. Prsts//ate flowers as in staminate, but ca-
lyx 0.6-0.9 mm long, the tube 0.2 mm long, the lobes 0.4-0.7 mm long
and wide, the margins opaque, entire; corolla, staminodes and pistil un-
known. Fruit dark purple at maturity, globose, 0.4—0.5 mm long, 0.4—0.6
mm diam., smooth, incospicuously pellucid punctate.

Distribution.—Upper Rio Santiago Valley and adjacent Serrania de Bagua,
Amazonas, 200—2,000 m, and Maynas Province, Loreto, Peru, along the
Rio Napo, at 120 m elevation.

Ecology and conservation status. —C ybianthus incognitus is mostly a ridgetop
species in the cloud forests of Amazonas Department, occurring infrequently
near the forest margin. The Upper Rio Santiago Valley and adjacent Serrania
de Bagua are known for their endemic species (Pipoly 1992b). However,
one surprising collection was noted at 120 m elevation, from Caserio de
Urcumirafio, in Maynas Province of Loreto Department, where vegetation
normally associated with much higher altitudes occurs on the tops of un-
dulating hills. Much more fieldwork will be required to better understand
forest dynamics at the western limits of the Peruvian Amazon with the foothills
of the Andes

Etymology.—The specific epithet refers to the fact that the plant was
misidentified even to family for nearly twenty years, and was finally identified
only when a flowering specimen was matched with the other fruiting specimens.
The densely spicate infructescences with numerous fruits were heretofore
unknown in the genus.

Paratypes. PERU. Amazonas: Prov. Bagua, 12 km E of La Peca, cloud forest, 1700 m,
20 Jun 1978 (fr), BP. Barbour 2405 (AMAZ, F, MO, USM), 29 Jun 1978 (fr), PR. Barbour
2567 (AMAZ, BRIT, F, MO, NY, US); Ca. 12-18 km E of La Peca in Serranifa de Bagua,
cloud forest, 1,800-1 95 )m, 14 Jun 1978 (fr), A. Gentry et al. 22859 (F, MO, USM), A.
Gentry et al. 22911 (F, MO, USM). Loreto: Rio Napo near entrance to Isla Inayuga, 20 Sep
1972 (fr), T. Croat 20528 (AMAZ, MO, USM); Caserio de Urcumirafio, Rio Napo, 2 hours
along trail from village to forest, 120 m, 8 Oct 1979 (stam. fl), C. Diaz & N. Jaramillo
1474 (AMAZ, BRIT, MO, USM)

Cybianthus incognitus is appears to be most closely related to Cybianthus
prieurit A. DC. of the Guianas, Venezuela and Brazil, because of the oblan-
ceolate, highly nitid coriaceous leaf blades and the filaments three times
longer than the anthers. However, Cybianthus incognitus is separated from
C. prieurii by its canaliculate petioles, dendroid and stellate tomentose branchlets,
sessile to subsessile flowers (spicate inflorescences), inconspicuously pellu-

Pipoty, Cybianthus in Ecuador and Peru 127

cid-punctate calyx lobes, ovate corolla lobes, globose pistillode and smooth
fruits. Cybianthus incognitus is unique within the subgenus by virtue of its
densely spicate inflorescences, dendroid and stellate tomentum of the branchlets
and inconspicuosly punctate calyx lobes.

32. Cybianthus minutiflorus Mez, Repert. Spec. Nov. Regni Veg. 3:102.
1906. Type: PERU. Loreto: near Rioja, W of Moyobamba, 800-900 m, 8 Sep 1904
(pist. fl, fr), A. Weberbaner 4699 (HOLOTYPE: B -destr.; fragment, F; LECTOTYPE, here
designated: F). Because the fragment at F contains floral and leaf material, and leave
no doubt as to the identity of the species, in the absence of other duplicates, it is
most appropriate to select this “clastotype” (a fragment taken with permission) as
the lectotype.

Tree to 3 m tall. Branchlets angulate, 4.5-6 mm diam., densely rufous
stellate-tomentose, glabrescent. Leaves alternate; blades membranaceous, widely
(rarely narrowly) oblanceolate, (14—)19-—28(—40) cm long, (4—)9-12(-15)
cm wide, apically acuminate, the acumen 1—1.5 cm long, basally cuneate,
midrib slightly raised above, prominently raised and densely rufous tomentulose
below, the secondary veins (9—)12—21 pairs, slightly raised above, promi-
nently raised and sparsely rufous tomentulose below, smooth and glabrous
above at maturity, sparsely rufous puberulent below, conspicuously pellu-
cid punctate, the margin entire, flat; petioles canaliculate, 1.7—2 cm long,
somewhat pulvinate, glabrous above, rufous tomentulose below, glabres-
cent. Staminate inflorescence a simple, erect raceme, 4.5—9 cm long, the ra-
chis densely rufous stellate-tomentose; floral bracts linear-lanceolate, 1—1.2
mm long, 0.2—0.3 mm wide, apically attenuate, densely tomentose above
and below, the margin entire; pedicels cylindrical, 0.6-0.9 mm long, densely
tomentose, persistent. Staminate flowers pale yellow; calyx carnose, cotyliform,
0.6-0.8 mm long, the tube ca. 0.1 mm long, the lobes widely triangular,
0.5—0.7 mm long, 0.7—-0.9 mm wide, apically acute, densely and promi-
nently orange punctate, sparsely rufous puberulent, the margin irregular,
opaque, somewhat crenulate, minutely glandular-ciliolate; corolla carnose,
subrotate, 1.2—1.4 mm long, the tube 0.3—0.4 mm long, the lobes widely
triangular, 0.9-1.1 mm long, 1.1-1.2 mm wide, apically obtuse, densely
and prominently orange punctate medially without, densely glandular-granulose
medially and above anther within, the margin opaque, glandular-granulose,
entire; stamens apparently sessile at corolla tube apically, the anthers widely
obcordate, 0.4—0.5 mm long, 0.7—0.9 mm wide, apically acute, basally cordate,
the thecae moderately yellow glandular-granulose, the connective promi-
nently red punctate; pistillode conic, 0.3-0.4 mm long, 0.1—0.2 mm diam.,
the stigma glandular-papillate. Pzsti//ate inflorescence as in staminate but (4—)
8-13 cm long; floral bracts 0.6—-1 mm long, 0.1—0.2 mm wide, apically
attenuate, densely tomentose above and below, the margin entire; pedicels
0.6—0.8 mm long, accrescent in fruit to 1.8 mm long. Pistillate flowers as in

128 Sipa 18(1)

staminate but green; calyx 0.8—1 mm long, the tube ca. 0.1 mm long, the
lobes 0.7—0.9 mm long, 0.9—1 mm wide; corolla 1-1.2 mm long, the tube
0.2—0.3 mm long, the lobes 0.8-1.0 mm long, 1—1.1 mm wide; staminodes
as in stamens but antherodes obcordate, 0.2—0.3 mm long, 0.2-0.3 mm
wide; pistil cylindrical, 0.6-1 mm long, 0.2-0.3 mm diam., the ovary an-
gular, 0.3—0.4 mm long, the style 0.3—-0.6 mm long, the stigma subcapitate,
4-lobed, the placenta cotyliform, ovules 2, naked. Fruit depressed-globose,
2.5—3.5 mm long, 4.5—5.5 mm diam., inconspicuously pellucid punctate,
the exocarp thin. Bisexual inflorescence in staminate but 2—3(—5) cm long,
the rachis moderately tomentose; floral bracts, 0.6-0.9 mm long, 0.1—-0.2
mm wide; pedicels 0.6-0.7 mm long. Bisexual flowers green; calyx 0.7—-8
mm long, the tube ca. 0.1 mm long, the lobes 0.6—0.7 mm long and wide;
corolla 1.3-1.4 mm long, the tube 0.2—-0.3 mm long, the lobes 1.1—1.2
mm long, 1—1.1 mm wide; stamens identical to pistillate staminodes, but
thecae full of pollen; pistil almost indistinguishable from pistillate flower
except the stigma subcapitate, 3—4-lobed, ovule 1, naked. Frwit (from bi-
sexual flower) unknown.

Distribution.—Endemic to the eastern slopes of the Andes and adjacent
Amazonian Hylaea of Peru and adjacent Bolivia, 100-1200 m

Ecology and conservation status. —Cybianthus minutiflorus occurs in primary
tall wet forest and premontane forest, on well drained white sands, known
as varillal in Peru. These pockets of sandstone often alternate with rolling
lateritic hills in the foothills of the eastern Andean slopes and adjacent Amazonia.
The lowland forests where Cybianthus minutiflorus is occurs are also notable
for their numerous lianas. Cybianthus minutiflorus is a rare species and should
be considered threatened.

Etymology.—The specific epithet refers to the extremely small flowers,
some of the smallest in the subgenus.

Local names and uses.—Peru: “taki kaspi” (Mayna Jivaro); leaves are boiled
in water and the decoction drunk to treat stomach ache.

Specimens examined. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Cerros de Putuim,
05° 03! 20"S. 78° 20! 23" W, 350 m, 15 Jun 1996 (stam. fl), R. Vasquez et al. 21187 (AMAZ,
MO). Prov. Condorcanqui, Dtto. El Cenepa, NE region of Marafion Drainage Basin, Rio
Cenepa, Comunidad Tutino, 04° 33'S, 78° 10' W, 750 m, 22 Nov 1993 (fr), R. Vasquez et
al, 18520 (BRIT, HUT, MO, USM). Cusco: Quipicanchi Prov., Camanti, Maniri, along
Rio Maniri and along the trail to Quebrada Garrote, 13° 71'S, 70° 45' W, 720 m, 8 Sep
1990 (bud), M. Timand 922 (CUZ, MO, USM). Huanuco: Rio Llulla Pichis watershed,
Cerros del Sira, 1,290 m, 17 Jul 1969 (fr), J. Wolfe 12346 (FE, NA, US). Loreto: Prov. Loreto,
Pampa Hermosa and vicinity, Rio Corrientes, | km S of junction with Rio Macusari, 03°
15'S, 75° 50' W, 160 m, 3-20 Dec 1985 (fr), W. Lewis et al. 10312 (BRIT, MO); Prov.
Maynas, Deto. Las Amazonas, Explornapo Camp, near Sucusari, along Rio Napo, 03° 20!
S, 72° 55' W, 100-140 m, 3 Mar 1991 (ster.), J. Pipoly et al. 14174 (MO, UNAP); Drto.
Iquitos, Allpahuayo (IIAP), Permanent inventory, 04° 10'S, 73° 30' W, 150 m, 25 Mar
1992 (ster.), R. Vasquez et al, 18163 (BRIT, MO, UNAP). Pasco: Oxapampa, Rd. in con-
struction between Oxapampa and Villa Rica, km 7, 10° 37'S, 75° 20! W, 2,100 m, 4 Jan

Pipoty, Cybianthus in Ecuador and Peru

5mm

Stet ope
ae ere

Ok Pe oe ee

ME A A oc

NE
See

¥

Ss

oe

m
caste
a re .

at

densely perpuncticulose leaf surface. C. P

habit, and calyx erose-dentate and gla
constricted corolla lobe b

Fic. 23. Cybianthus huampamiensis Pipoly. A. Habit, showing pulvinate petioles. B. Abaxial,
: re £ ‘ 4 | : oe

ndular-ciliate. D. Staminate flower, showing abruptly
ase. E. Fruit and calyx. FE Branchlet apex. G. Pistillate flower,
showing constricted corolla lobe base. A-B, drawn from Kayap 783. C—D, drawn from Kayap
982. E-G, drawn from Kayap 933. Figure drawn by Linda Ellis.

130 Sipa 18(1)

1984 (bisex. fl), R. Foster et al. 7817 (F, MO, USM); Oxapampa, Pichis Valley, San Matias
Ridge, 10-12 km SW of Puerto Bermiidez, above Sta. Rosa de Chivis, trail to Loma Linda,
10° 20'S, 75° 00' W, 1,000 m, 29 Sep 1982 (fr), R. Foster 8624 (F, MO, USM), (stam. fl),
R. Foster 8981 (BRIT, MO, F, NY, US, USM). San Martin: Chazuta, Rio Huallaga, 260 m,
Mar 1935 (stam. fl), G. K/vg 3981 (F, MO, 5S, US); Prov. Mariscal Caceres, Dtto. Tocache
Nuevo, Isla de Pucunchu, right bank of Rio Huallaga, 3 Apr 1971 (fr), J. Shunbe 4779 (F,
MO, NY, US); W of bridge, 700-800 m, 16 Dec 1971 (fr), J. Schunke 5737 (F, NY, MO,
US). Ucayali: Prov. Coronel Portillo, Plantacién Margarita, near Loreto border, 1,500—
1,600 m, 14 Aug 1946 (fr), R. Ferreyra 1040 (US, USM). BOLIVIA. Pando: Prov. Madre
de Dios, Mobil Oil Site, 12° 10'S, 67° 15' W, 170 m, 20-25 Aug 1992 (fr), T: Killeen
4449 (BRIT, LPB, MO).

Cybianthus minutiflorus is most closely related to C. granulosus Pipoly by
its densely rufous furfuraceous-lepidote branchlets, erect, carnose, densely
and prominently black punctate perianth, and sessile anthers. However, the
flat petioles, subacuminate leaf apices, short pedicels, opaque, crenulate and
epunctate margins of the calyx lobes, and emarginate anthers easily distin-
guish Cybianthus minutiflorus.

33. Cybianthus huampamiensis Pipoly, sp. nov. (Fig. 23). Type: PERU. AMAZONas:
uebrada chigkan entsa, Rio Cenepa, 300 m, 9 Jun 1973 (stam. fl), E. Ancuash 588
(HOLOTYPE: MO; isotypes: NY, USM).

Propter folia elliptica lanceolata vel ob! lata, longipetiolat i] le vel inequilaterale

fl graciles fore erectes deminutosque,

secus margines irregulares, rhachides 1
necnon frutos minores, ad aspectu primo intuito C. resimoso arcte similans sed ab ea laminis
membranaceis (non tenuiter coriaceis), utrinque sordidis (nec nitidis) subter manifeste

prominenteque atro-perpuncticulosis (nec epunctatis), petiolis (1.5—)2—2.5 (non 0.5—1.4)
cm longis, lobis calycinis translucentibus (non opacis) acuminatis (nec rotundatis), lobis
corollinis extus glandulari-granulosis (non glabris) acutis vel rotundatis (nec obtusis vel
emarginatis) denique fructibus luteis (non atris) permanifeste distinguitur.

Tree to 3(—6) m tall. Branchlets subterete to angulate, (3.5—)4—5 mm diam.,
densely rufous tomentose, glabrescent. Leaves pseudoverticillate; blades
membranaceous, elliptic lanceolate or rarely, oblanceolate, (12—)16-25(—
31) cm long, (5.2—)7—9(—12) cm wide, apically long-acuminate, basally acute,
not decurrent on the petiole, dull green above, pallid below, midrib im-
pressed above, prominently raised below, the secondary veins 12—18 pairs,
prominently raised below, glabrous above rufous puberulent below along
the veins, prominently and densely perpuncticulose below, the margin en-
tire, irregular, flat; petiole slightly canaliculate distally or flat, 2—2.5 mm
long, thick and pulvinate, densely rufous puberulent at first, glabrescent.
Staminate inflorescence a lax, simple, subspicate raceme, (S—)10—(18) cm long,
sparsely rufous puberulent; floral bracts linear-lanceolate, 1-1.3 mm long,
apically attenuate, densely rufous publerulent abaxially, the margin erose,
persistent; pedicels cylindrical 0.8—0.9 mm long, densely rufous pubescent,
glabrescent. Staminate flowers yellowish to orange; calyx membranaceous,
cotyliform, translucent, 1.2-1.4 mm long, the tube ca. 0.2 mm long, the

Prpoty, Cybianthus in Ecuador and Peru 131

lobes ovate, 1—-1.2 mm long, 0.6—0.8 mm wide, apically subacuminate, densely
and prominently orange punctate, glabrous, the margin hyaline, prominently
erose-dentate, sparsely glandular-ciliate; corolla chartaceous, subrotate, 2.0—
2.4 mm long, the tube 0.4—0.5 mm long, the lobes ovate, 1.6-1.9 mm long,
1.2-1.4 mm wide, apically acute to rounded, sparsely glandular-granulose
without and densely so throughout within, densely and prominently or-
ange punctate, flat, the margin scarious, erose-denticulate and glandular-
granulose; anthers apparently sessile at junction of corolla tube and lobe,
very widely ovate, 0.4—-0.5 mm long, 0.6-0.8 mm wide, apically obtuse to
rounded, dehiscent by apical confluent pores extending ca. 2/3 length of
anther, the connective epunctate, densely rubiginous glandular-papillate dorsally;
pistillode conic, ca. 0.2—0.3 mm long and diam., hollow, glandular-papil-
late. Pistillate inflorescence as in staminate but (S—)7—9(—-14) cm long, densely
rufous puberulent at first, glabrescent; pedicel 0.4—0.5 mm long. Pzistillate
flowers as in staminate but yellowish to orange; calyx 1—-1.2 mm long, the
tube ca. 0.2 mm long, the lobes 0.8—1 mm long, 0.4—0.5 mm wide; corolla
1.8—2.2 mm long, the tube 0.3—0.4 mm long, the lobes 1.5—1.7 mm long,
1.1-1.4 mm wide; staminodes as in stamens but antherodes 0.3—0.4 mm
long, 0.5—0.6 mm wide, apically rounded to acute; pistil conic, ca. 1.3 mm
long, the ovary 0.5—0.6 mm long 0.6—0.8 mm diam., translucent glandu-
lar lepidote, the style short, to 0.2 mm long, the stigma 4-lobate, the lobes
distally curved, glandular-papillate. Frvit yellow, subglobose, 4—6 mm long,
5—8 mm diam. inconspicuously pellucid punctate.

Distribution.—Endemic to the Rio Marafion, Rio Cenepa and Rio Santiago
drainage basins in the northwest corner of the Department of Amazonas,
Peru, 200—550(—1,850) m.

Ecology and conservation status. —C ybianthus huampamiensis occurs in the foothills
of the premontane and lowland wet forest of the most underexplored area
of the Peruvian Hylaea/Andean interface. Given that the region is a border
area, and therefore, a priority for development, this species should be con-
sidered.

Etymology.—The epithet describes the place where the many of the col-
lections were made, the Haumpami area of the Rio Cenepa Drainage Basin.

Paratypes. PERU. Amazonas: Prov. Bagua; ca. 12—18 trail km E of La Peca, Serranfa
de Bagua, 1,800-1,950 m, 14 Jun 1978 (fr), A. Gentry et al. 22859 (F, MO, USM); Rio
Cenepa, Quebrada tujushik entsa, 330 m, 18 Apr 1973 (fr), E. Ancuash 274 (MO, USM);
along Rio Cenepa, 350 m, 3 May 1973 (fr), E. Ancuash 303 (MO, USM); Rio ae Quebrada
Idayua entsa, 400 m, 16 May 1973 (fr), E. Ancuash 392 (BRIT, F, LL-TE O, US, USM);
Rio Cenepa eee Wampusik entsa, 5} Aug 1974 (stam. fl), E. ee FS ((MO-2
sheets USM); R o Cenepa, Quebrada Cikan Inci, 250-330 m, | Jan 1973 (stam. fl), B.
Berlin 779 (MO, USM): 10 km N of Quebrada Huampami, 200-250 m, 24 Jul 1974 (stam.
fl), B. Berlin 1760 (MO, USM); Quebrada Chigkui Shinuki Cenepa, 250 m, 11 Apr 1973
(fr), R. Kayap 618 (MO, USM), 23 May 1973 (stam. fl), R. Kayap 783 (MO, NY, USM);

Stipa 18(1)

=
No

BF hese ye.

Fic. 24. Cybianthus granulosus Pipoly. A. Habit. B. Branchlet apex. C. Abaxial leaf surface,
showing minutely rufous lepidote indumentum. D. Staminate flower. E. Staminate corolla.

F. Staminate calyx. A-C, drawn from holotype. D-F, drawn from V. Huwashikat 1221.

Figure drawn by Linda Ellis.

Prpory, Cybianthus in Ecuador and Peru 135
Quebrada Etseketai, Rio Cenepa, 250 m, 31 May 1973 (fr), R. Kayap 856 (MO, USM);
Quebrada Wampushik entsa, 330 m, 13 Jun 1973 (pist. fl, fr), R. Kayap 933 (F, LL-TEX,
MO, NY, USM); Huampami, Rio Cenepa, 200 m, 15 Jun 1973 (stam. fl), R. Kayap 982
(LL-TEX, MO, NY, USM); Rio Cenepa, vicinity Huampami, ca. 5 km E of Chavez Valdivia,
04° 30'S, 78° 30! W, 200-250 m, 31 Jul. 1978 (fr), A. Kujikat 50 (LL-TEX, MO, NY,
USM), 12 Aug. 1978 (fr), A. Kujikat 291 (MO, NY, USM), 15 Aug. 1978 (fr), A. Kujikat
395 (MO, USM); Mouth of Rio Santiago, without date (stam. fl), G. Tessmann 4439 (NY);
Rio Santiago Me 03° 50'S, 77° 40' W, Quebrada Caterpiza, 2—3 km from Caterpiza,
as m, 28 Dec 1979 (stam. fl), 8. Tunquf 488 (MO, NY, USM), Deto. Imaza, Quebrada
za, | hour aie from Comunidad de Cunchim, 370 m, 21 Jul 1994 (fr), C. Draz et al.
pe 30 (BRIT, HUT, MO, USM); Rio Marafion Drainage Basin, Comunidad de Yamayakart,
Rio Marafion, 04° 55'S, 78° 19' W, 600 m, 28 Jan 1995 (fr), E. Rodriguez 283 (BRIT,
HUT, MO, USM), Quebrada Kusu-Chapi, 04° 55'S, 78° 19' W, 550 m, Feb 1995 (stam.
fl), R. Vague et al. 20045 (BRIT, HUT, MO), 320 m, 11 Mar 1996 (stam. fl), N. Jaramillo
et al. 1351 (BRIT, HUT, MO).

Cybianthus huampamiensis may at once be distinguished from all other species
of the subgenus by its translucent calyx, externally glandular-granulose corolla
and yellow fruit. This species is one of many endemic taxa known from this
most underexplored area at the junction of the eastern Andean slopes with
the Amazon Basin in northern Peru.

34. Cybianthus granulosus Pipoly, sp. nov. (Fig. 24). Tyrer: PERU. Amazonas:
Rio Santiago Valley, 03° 50'S, 77° 40! W, Quebrada Caterpiza, 2—3 km from Caterpiza
settlement, primary forest, 200 m, treelet 2 m tall, fls. brownish-green, 28 Nov 1979
(stam. fl), S. Tungué 161 (HoLoTYPE: MO; Isorype: NY).

uoad folia alterna chartacea ad apices acuminata ad pas cuneataque, ammlos dense

rufo- furfuraceo-lepidotos, flores GLEE tos, Carnosos, t an nthe ras

iles, ad C. minutifloro valde affinis sed ab ea petiolis conaliculaeis (non planis) laminis
abrupte largo- sos sub- Vac Men ial, ao 15-2. " nee 0.5— 1) mom nes lobis calyeinis

lobis eaiellinis secus crakear erosis (nec integerrimis) amihienis acutis (nec sennesiiatis)
facile cognoscitur.

Treelet to 3 m tall. Branchlets angulate, 2-4 mm diam., densely rufous
furfuraceous lepidote. Leaves alternate; blades chartaceous, elliptic to nar-
rowly oblanceolate, (9Q—)11—15(—18) cm long, (3—)4—5(—6) cm wide, apically
abruptly acuminate, basally cuneate, densely rufous lepidote (appearing granulose)
above and below at first, glabrate above, somewhat persistent below, mid-
rib impressed above, prominently raised below, the secondary veins 8-12
pairs, impressed above, prominently raised below, inconspicuously pellu-
cid punctate, the margin entire; petioles canaliculate, 1.5—1.8 cm long, glabrescent
above, densely lepidote below. Staminate inflorescence a simple raceme, (4—
6—-9(-10)cm long, the rachis and pedicels densely rufous furfuraceous lepi-
dote; floral bracts carnose, linear lanceolate, 0.8—1.2 mm long, apically long-
attenuate, the margin entire, densely lepidote; pedicels cylindrical, 1.9—
2.2 mm long. Staminate flowers erect, 4-merous, carnose, subrotate, brownish-green;

134 Sipa 18(1)

calyx 0.8-1.1 mm long, the tube 0.1—0.2 mm long, the lobes ovate, 0.7—
0.9 mm long, 0.8—0.8 mm wide, apically acute, abruptly constricted ba-
sally, densely and prominently black punctate, moderately rufous lepidote,
glabrescent, the margin hyaline, conspicuously black punctate-lineate, ir-
regularly serrate, glabrous; corolla subrotate, 1.6—1.8 mm long, the tube
0.3—0.4 mm long, square, glabrous, the lobes suborbicular, 1.3—1.4 mm
long and wide, emarginate apically, abruptly constricted basally, densely
and prominently black punctate, sparsely rufous lepidote without, glabrescent,
densely glandular-granulose throughout within, the margin erose; anthers
sessile at apex of corolla tube, thus appearing epipetalous, the tube ()..3—0.4
mm long, glabrous, the anthers widely cuadrate, 0.3—0.4 mm long, 0.6—
0.7 mm wide, apically acute, basally truncate, dehiscent by large subapi-
cal, ovate pores, the pores not confluent, extending ca. 3/4 anther length,
the connectives densely and prominently red or black punctate; pistillode
conic, 0.2—-0.3 mm long, sparsely glandular-lepidote. Pistillate inflorescence
unknown. Frait unknown.

Distribution.—Endemic to the upper Rfo Santiago Valley, in the Depart-
ment of Amazonas, Peru, 180—200 m.

Ecology and conservation status. —Cybianthus granulosus inhabits wet e
forests above the Rio Santiago valley, which together with the Rio Cenepa,
comprise a region now known to be host to a number of endemic species.
Given that it is most likely endemic, and not at all well-known, its conser-
vation status cannot be determined at this time.

Etymology.—The specific epithet refers co the densely rufous lepidote
tomentum, which appears granulose when examined superficially

Paratyres. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Comunidad Aguaruna Putuim,

xo Yamayakat, Zonas Altas de Putuim, “Campou,” 700 m, 18 Jan 1996 (fr), C. Daz et

a 76494 (BRIT, HUT, MO, USM); Rio Sasi 3 km from Caterpiza, 180 m, 12 Nov

1979 (stam. fl), V. Haashikat 1221 (MO, USM); Valle del Rio Santiago, 65 km N of Pinglo,

Quebrada Caterpiza, 2—3 km from nea 200 m, 19 Sep 1977 (bud), V. Huashikat 677
(MO, USM), 28 Nov 1979 (stam. fl), V. Fuging 18 22 (MO, USM).

Cybianthus granulosus is most closely related to C. minutiflorus Mez by virtue
of its alternate, chartaceous leaf blades with acuminate apices and cuneate
bases, densely rufous furfuraceous-lepdidote branchles, flowers with carnose
texture, densely and prominently black punctate, and sessile anthers. However,
Cybranthus granulosus is easily separated from C. minutiflorus by the abruptly
long-acuminate leaf apices, canaliculate petioles, the hyaline, serrate, and
punctate-lineate calyx margins, the erose corolla margins, acute anthers, and
pedicels 1.5—2.5 mm long.

35. Cybianthus flavovirens Pipoly, sp. nov. (Fig. 25). Type: PERU. SAN Martin:

rov. Mariscal Caceres, Dtto. Tocaché Nuevo, Palo Blanco near Fundo de Manuel

Pipoty, Cybianthus in Ecuador and Peru 135

at

}
}.
x
4
h
t
§
ae

A ate

Fic. 25. Cybianthus flavovirens Pipoly. A. Habit, showing angulate branchlets and gradually
tapering petioles. B. Abaxial leaf surface, showing sparse, rufous stellate tomentum. C.
Section of inflorescence, showing nodding flowers, widely deltate calyx lobes and corolla
with lobes constricted basally, prominent veins, and pusticulate surface. D. Branchlet, showing
rufous stellate tomentum. A—D, drawn from holotype, by Linda Ellis.

136 SIDA 18(1)

Aranjo, 700-800 m, 1 Mar — fl), J. Schunke 10883 (HOLOTYPE: MO; ISOTYPES:
AMAZ, BRIT, EF, TEX, US, USM).

Propter ramulos angulatos, petiolos pracate BE USEAtOSs aor racemosam, flores
nutantes, antheras sessilia, C. veneznelam eda
(non carnosis) acutis (nec rotundatis) secus marginis cola Ge Space: antheris obtusis

vel ee (non Eeuncatls), pot sgue sepatans (ee sonBucacens), denique lobis ene

praeclare distat. .

Tree to 4 m tall. Branchlers lightly angulate, 2.5—3.5 mm diam., densely
rufous stellate-tomentose. Leaves alternate; blades chartaceous, elliptic, (12.5—)
19—30(—32) cm long, (4—)6—10 cm wide, apically subacuminate to acumi-
nate, the acumen 0.5—2 cm long, basally attenuate, decurrent on the distal
end of the petiole, midrib somewhat elevated but canaliculate above, not
decurrent on petiole, prominently raised and densely rufous tomenose be-
low, the secondary veins 10—16 pairs, dull green above, pallid green below,
pellucid punctate above and below, sparsely rufous stellate-puberulent be-
low, the margin entire, opaque, regular, flat; petioles semiterete (1.6—)2—
2.5 cm, flat above, tapered, slightly thickend basally, not pulvinate, gla-
brous above, sparsely rufous pubescent below, glabrescent. S/aminate inflorescence
a lax raceme (7.5—)13—19(—40) cm long, the rachis and pedicels cana
rufous tomentellous, glabrescent; floral bracts linear-lanceolate, 1.1—1.5 m
long, 0.1-0.2 mm wide, apically attenuate, densely rufous eeeae ioe
below, glabrescent, the margin entire; pedicels cylindrical, 2.5—3.5 mm long
at anthesis, erect in bud, nodding in anthesis. Staminate flowers 4-merous,
membranaceous, nodding, bright yellow-green; calyx cotyliform, 1.5—
mm long, the tube 0.6—0.8 mm long, the lobes widely deltate to triangu-
lar, 1.1-1.3 mm long and wide, apically acute, epunctate, medially thick-
ened, sparsely rufous puberulent, glabrescent, the margin hyaline, € epunctate,
entire, minutely glandular-ciliolate; corolla subrotate, 2.8—3 mm long, the
tube 0.5—0.8 mm long, glabrous without, densely glandular-granulos within,
the lobes suborbicular to oblate, 1.5—2.2 mm long, 2.2—2.6 mm wide, apically

truncate to slightly emarginate, one bacall ly, cache the three
veins conspicuous, glabrous without, prominently pusticulate and sparsely
glandular-granulose above but densely so toward base within, the margin
opaque, sparsely glandular-granulose, entire; stamens apparently sessile at
junction of corolla lobes and tube, the anthers sessile, very widely ovate-
obcordate, 0.4—0.6 mm long, 0.5—0.8 mm wide, apically obtuse to sub-
acute, basally cordate, the pores widely ovate, extending 1/2—3/4 anther
length, ser (not confluent), the connective epunctate, minutely red glandular-

papillate; pistillode costate, subglobose, 0.8—1 mm long, 0.6—0.7 mm wide,

eon

hollow, the stigma capitate, densely yellow glandular-papilalte. Pisri//ate
inflorescence: unknown. Fruit unknown.

Pipoty, Cybianthus in Ecuador and Peru To?

Distribution.—Endemic to the junction of the westernmost Amazon Ba-
sin (Hylaea) with the foothills of the Peruvian Andes, from 100-800 m elevation.

Ecology and conservation status. —Cybianthus flavovirens occurs in lowland
moist forests on serra firme. These forests are drier than sites with the same
general physiognomy farther to the north in Amazonas and Loreto. The few
sporadic collections may be the result of underexploration rather than rar-
ity, but owing to increasing pressure to cut forests for farming, this species
should be considered threatened.

Etymology.—The epithet refers to the bright yellow-green corolla, a unique
feature in the subgenus.

Paratypes. PERU. Hudnuco: Villa Isabel, Rio Cuchara, 20 Sep 1961 (stam. fl), J. Schunke
5667 (F, K, MO, US, USM). Loreto: Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929 (stam
fl), G. Klug 285 (EF, US), (stam. fl), G. K/vg 367 (EF, US). Pasco: Prov. Oxapampa, Palcazu
Valley, Cabeza de Mono, 5—6 km W of Iscosacfn, 10° 12'S, 75° 14' W, 14-15 Apr 1983
(stam. fl), D. Smith 3709 (MO, US, ee Prov. Pasco, Palcazu Valley, Sela Central, Cerro
de Pasco, Proyecto Especial Pichis—Palcazu, IND, 09° 50! S, 68° 00’W, 300-600 m, 13
Oct 1987 (stam. fl), G. ae ey et al, 2996 (BRIT, MO, USM).

Cybianthus flavovirens isa member of a complex of taxa related to C. venezuelanus
Mez, as evidenced by the angulate branches, gradually tapered petioles, nodding
flowers, and sessile anthers. However, C. flavovirens is easily recognized by
its membranaceous perianth, acute calyx lobes with opaque margins, pusticulate,
bright yellow-green corolla lobes with truncate or emarginate apices, and
obtuse or subacute anthers with separate pores. This species has been con-
fused with Cybianthus cyclopetalus. However, Cybianthus flavovirens may eas-
ily be separated from that species by its angulate branchlets, racemose
inflorescences, large, yellow flowers, and deltate to triangular calyx lobes.

36. Slants venezuelanus Mez in Engl., Pflanzenr. IV. 236(Heft 9):221.
1902 E: VENEZUELA. Lara: Barquisimeto, San Felipe, 600 m, Jun 1846 (stam.
fl), N. ey & L. Schlim 678 (LECTOTYPE, here Be cea fe BOIS; 1soLECTOTYPES:

, sAa; LD; PW)

Peckia purpurea Rusby, Bull. New York Bot. Gard. 4:405. 1907. Type: BOLIVIA. With-
out locality data, (stam. fl), A.M. Bang 2048 (HoLoTYPE: NY).

Cybianthus egensis Mez in Engl., Pflanzenr. IV. 236(Heft 9):222. 1902. syn. nov. Ty
BRAZIL. AMazonas: Near Ega [Teffel, Sep 1831 (stam. fl), E. Poeppig 2567 Pee
TYPE: W, F Neg. 31997; IsorypeE: W

Tae. browntt eigen. Bull. Torrey Bot. Club 53:293. 1926. syn nov. Type: GUYANA

H GUIANA}. Tumatumari, 18 Jun—8 Jul 1921 (stam. fl), H. Gleason 159
aa PE: NY; ISOTYPE: K).

Tree to 5 m tall. Branchlets angulate, (2.5—)3.5—5 mm diam., densely ru-
fous tomentose. Leaves pseudoverticillate; blades chartaceous, oblanceolate
to elliptic, (10—)17—27(—34) cm long, (3—)6—9(—11) cm wide, apically acute
to acuminate, basally acute, slightly decurrent on the petiole, midrib slightly

138 Sipa 18(1)

raised above, prominently raised below, the secondary veins 10—13 pairs,
rufous puberulent and smooth above, glabrescent, sparsely rufous puberu-
lent below, concentrated along the midrib and secondary veins, inconspicuously
pellucid punctate, the margin flat, entire; petioles slightly canaliculate, (15—)
20—25(—30) mm long, tapered, densely and minutely stellate rufous stel-
late puberulent. Staminate inflorescence: an erect, simple raceme, (7.5—)10.5—
18(—23) cm long, sparsely rufous stellate puberulent; peduncle 1—3 cm long;
floral bracts coriaceous, lanceolate, 0.5—0.7 mm long, 0.2—0.3 mm wide,
apically subulate, densely rufous stellate puberulent, the margin entire; pedicels
cylindrical, 2.1-2.7 mm long, sparsely rufous stellate puberulent, glabres-
cent. Staminate flowers 4-merous, carnose, nodding, green; calyx cotyliform,
0.9-1.1 mm long, the tube ca. 0.1 mm long, the lobes widely ovate, 0.8—
| mm long, 0.9—1 mm wide, apically rounded, densely rufous stellate pu-
berulent, sparsely and prominently black punctate, the margin opaque, coarsely
serrulate, densely glandular-ciliolate; corolla carnose, subrotate, 1.6—1.8 mm
long, the tube quadrate, 0.5—0.6 mm long, glabrous, the lobes widely tri-
angular, 1.1-1.3 mm long, 1.5—1.7 mm wide, reflexed in anthesis, apically
acute, dorsally recurved, prominently and densely black punctate and gla-
brous without, densely glandular granulose throughout within, the mar-
gin slightly revolute, densely glandular-granulose; staminal tube inconspicuous,
hyaline, membranous, 0.5—0.6 mm long, adnate to corolla tube, anthers
apparently sessile, cuadrate, 0.3—0.4 mm long, 0.5—0.6 mm wide, apically
truncate, basally truncate, leaning proximally at anthesis, the connective
prominently punctate dorsally, rufous glandular-papillate apically; pistillode
obsolete or conical, 0.4—0.5 mm long, 0.1—0.2 mm wide, hollow, glabrous.
Pistillate inflorescence as 1n staminate but (4.5—)8.5—10(-13) cm long; peduncle

1—2.5 cm long; floral bracts 0.5—0.7 mm long, 0.2—0.3 mm wide; pedicels
0.9-1.2 mm long. Prsti/late flowers as in staminate but calyx 0.8-1.2 mm
long, the tube ca. 0.2 mm long, the lobes 0.6—0.8 mm long, 0.9-1.1 mm
wide; corolla 1.2—1.4 mm long, the tube 0.4—0.5 mm long, the lobes 0.7—
0.9 mm long, 1.1—1.2 mm wide; staminodial as in staminal tube, 0.4—0.5
mm long, adnate to corolla tube, the antherodes 0.2—0.3 mm long, 0.4—
0.5 mm wide; piscil obcurbinate, 0.5—0.6 mm long, 0.3—0.4 mm wide, glabrous,
the ovules 2—3, partially imbedded in the placenta. Fruit globose, 5—7(—9)
mm diam., black at maturity, the exocarp thick, juicy.

Distribution.—In moist forests rimming the Amazon Basin, from Guyana
through Venezuela to the Andes of Colombia southward to Bolivia, 100—
1,000(—2,000) m. It is also known from the Chocé Floristic Province of Colombia,
and may be expected in that corresponding region of Ecuador. The Boliv-
ian specimens cited below represent new distribution records.

Ecology and conservation status. —C ybianthus venezuelanus is common in lowland
and premontane moist forests, and occasionally in premontane pluvial for-

Pipoty, Cybianthus in Ecuador and Peru 139

ests. It is locally common and appears to do well in light gaps and forest
margins, thus, it should not be considered threatened.

Etymology.—The epithet refers to the the country from which the type
was collected, Venezuela.

Representative specimens examined. ECUADOR. Zamora-Chinchipe: Nangaritza Canton,
Pachicutza, Camino al Hito, Cordillera del Céndor, 04° 07'S, 78° 37! W, 1,000—1,100 m,

19 Oct 1991 (fr), WY Pai et al. 8313 (BRIT, MO, QCNE). PERU. Hence: Prov.
Leoncio Prado, Dtto. Rupa Rupa, Ynti, Rd. to Rfo Rondos, 750 m, 24 Mar 1972 (stam.
fl), J. Schunke 5308 (AMAZ, F, G, MO, NY, US, USM). Loreto: Prov Maynas, Mishuyacu,
near Iquitos, 100 m, Oct—Nov 1929 (stam. fl), G. Klug 94 (F, NY, US); Pefia Negra, near
Iquitos, 100 m, 20 Oct 1979 (bud), F Ayala 2102 (AMAZ, MO, Ree Quebrada Orején,
aes oe Oct 1980 (fr), E Ayala et al. 2814 (AMAZ, MO, US, USM); ae be-
tween | Moy oe mba, 600—1,200 m, Aug—Sep 1933 (stam. fl), G. Klug 3188
(F, G, MO, NY, S, US). Madre de Dios: Prov. Tambopata, Tambopata Wildlife sre
30 km S of Puerto ene 12° 15'S, 69° 17' W, 260 m, 14 Nov 1984 (stam. fl), H
ne et al. 179 (MO, US, USM); Tambopata Tourist Camp at junction of Rios Tambopata
Torre, 12° 49'S, 69° 43' W, 280 m, 22 Jul 1985 (ster.), A. Gentry et al. 51083
con MO, USM), 27 Jul 1985 (ster.), A. Gentry et al. 51400 eee MO, US, USM). BOLIVIA.
Santa Cruz: Prov. Florida, Parque Nacional Amboro, Sta. a de Lima, 5-8 km N of
Cerca a La Cumbre, on path to La Playa, 17° 49.5'S, 64° 16' ey Syren m, 3—5 May
1993 (fr), 1. Vargas et al. 2248 (BRIT, MO, USZ)

a

The perianth of the lectotype of Cybianthus venezuelanus, with dense and
prominent black punctations, the truncate sessile anthers, large oblanceolate
chartaceous leaves, and short pedicels match the collections cited here. Cybianthus
venezuelanus, as treated here, is a polymorphic ochlospecies with many re-
gional variants. The type of Cybzanthus brownii Gleason varies only in its
shorter pedicels aus eo ee colons lobes. Likewise, the type of Cybianthus
egensis Mez, in bud, gly shorter pedicels and corollas. Cybianthus
venezuelanus iS oe peer by the angulate branchlets, pseudoverticillate
leaves, rounded calyx lobes with opaque margins, and truncate anthers with
confluent pores. Its distribution, which rims the Amazon Basin, is unusual
in the Myrsinaceae but is common in other families, such as the Piperaceae
(R. Callejas, pers. comm.).

ore hea grandezii Pipoly, sp. nov. (Fig. 26). Typr: PERU. Loreto:
aynas, Quebrada Yanayacu, entering from Aucayo, 25 Aug 1990 (stam. fl),
C. CHE S. Vasquez & M. Flores 1824 (HOLOTYPE: MO; tsorypes: AMAZ, US).

J i]
Quoad folia magna chartacea laminares raeciare utrinque

conspicua ad bases gradatim descrescentiaque necnon ase pulvinatos C. jensoni valde
affinis sed ab ea laminis ad bases obtusis auriculatisque (non cuneatis) manifeste desuper
perpuncticulosis (nec epunctatis) desuper nitidus (nec sordidis) nerviis secundariis 18-24
(non 24—30) -jugis, 12—13.5 (non 6.5—9) cm latis, petiolis 0.5—1.4 (non 2.1—5) cm longis,
pedicellis 0.3—0.5 (non 0.8-1.4) mm longis, bracteis florinis pedicellis 3 (non 6—7)-plo
longiores statim cognoscitur.

Shrub to 2 m tall. Branchlets terete, ca. 10 mm diam., rufous-lepidote.

140 Sipa 18(1)

fl. ro
y,

cS
Fic. 26. Cybianthus grandezii Pipoly. A. Habit g
leaf bases. B. Adaxial, prominently perpuncticulose surface. C. Abaxial, minutely rufous
lepidote surface. D. Portion of inflorescence. E. Pistillate calyx, showing pustulate, black
punctate lobes with hyaline, erose-serrulate margins. F. Pistillate corolla, showing pustu-
late abaxial surface. G. Branchlet apex. A-G, drawn from holotype by Linda Ellis.

] : | 1] ] } - ]

Pipoty, Cybianthus in Ecuador and Peru 141

Leaves alternate; blades chartaceous, oblanceolate, 34—36 cm long, 12—13.2
cm wide, apically subacuminate, basally truncate and appearing auriculate,
nitid above, pallid below, glabrous, and prominently black perpuncticulose
above, sparsely and minutely rufous-lepidote below, the midrib raised and
canaliculate above, prominently raised, black punctate-lineate and densely
rufous-lepidote below, the secondary veins 18-24 pairs, loop-connected
submarginally, the margin entire; petioles canaliculate, (0.5—)1-1.4 mm
long, 0.4—0.6 mm diam., pulvinate, densely lepidote. Staminate inflorescence
unknown. Pisti/late inflorescence an erect, straight simple raceme, (8.5—)11—
14 cm long; peduncle 1—3 cm long; the rachis green, densely red glandu-
lar-papillate, black punctate-lineate; floral bracts membranaceous, linear-
lanceolate, 1.8—2 mm long, densely rufous lepidote; pedicels obconic, 0.5—1
mm long, densely glandular-papillate. Pisti//ate flowers chartaceous, creamish-
white; calyx cotyliform, 1.8-2 mm long, the tube ca. 0.5 mm long, the
lobes widely ovate, 1.3-1.5 mm long, 1.1-1.2 mm wide, apically acute,
densely and prominently black punctate, prominently translucent pustu-
late, the margin hyaline, irregular, erose-serrulate, epunctate, glabrous; corolla
subrotate, 2.2—2.4 mm long, the tube ca. 0.5 mm long, the lobes very widely
ovate 1.7-1.9 mm long and wide, apically rounded, densely and promi-
nently black punctate and translucent pustulate without, glandular-granulose
throughout and prominently black punctate within, the margin hyaline,
epunctate, minutely erose- crenulate, glabrous; staminodes 1.1—1.2 mm long,
staminal tube conspicuous, carnose, 0.4—0.5 mm long, elobate, densely glandular-
papillate, the filaments terete, proximally curved, 0.2—0.3 mm long, the
sterile anthers ovate, 0.5—0.6 mm long, and wide, the apically apiculate,
dehiscent by terminal confluent pores ca. 3/4 length, the connective prominently
black punctate; pistil obturbinate, 1.5—1.6 mm long, 0.5—0.6 mm diam.,
densely and prominently black punctate, translucent glandular-lepidote,
the ovary 0.5—0.6 mm long, the style 0.8-0.9 mm long, the stigma bi-
lobed, the lobes to 0.1 mm long, distally curved, the placenta subglobose,
with 2—4 ovules partially embedded. Fruit globose, 8-10 mm long and in
diam., the exocarp thick, juicy, purple-black at maturity.

Distribution. —Known only from Maynas Province, Dept. of Loreto, Peru,
at up to 125 m elevation; presumably endemic.

Ecology and conservation status.—Cybianthus grandezii is restricted to pri-
mary lowland most upland terra firme forest over red lateritic clays. Label
data indicate it is rare, and given the valuable timber present in that forest
type, this species should be considered threatened.

Etymology.—It is indeed a pleasure to dedicate this species to Bidl. César Grandez,
professor of biology at the Universidad Nacional de la Amazonia Peruana
(UNAP), Iquitos, and authority on the systematics of Peruvian Flacourtiaceae.
César is an ardent field worker, an excellent teacher and herbarium curator.

142 Stipa 18(1)

Pal : PERU. Loreto: Prov. Maynas, Dtto. Fernando Lores, Panguana, 125 m, 6
Aug | a te), S. McDaniel & M. Rimachi 31219 (MO, IBE).

Cyhianthus grandezit is closely related to C. jensonii Pipoly, but is easily
recognized by its subsessile leaves with truncate bases. The perpuncticulose
and nitid adaxial leaf surface, and secondary veins raised prominently above
and below are also distinctive.

38. Cybianthus jensonii Pipoly, sp. nov. (Fig. 27). Type: PERU. Loreto: Prov.
Alto oles Andoas, Rio Pastaza near Ecuadorean border, 02° 48'S, 76° 28'
210 m, 14 Aug 1980 (fo), . Gentry, R. Vasquez & N. Jaramillo 29700 (soueren:
MO; isotypes: AMAZ, NY, USM).
Quoad folia magna hartacea nervos | larios tertias 1osque praeclare utrinque
conspicua ad bases gradati | poclesdus pulvinatos C. wanes ae Hanis sed

d
sat }

perpuncticulosis) desuper sordidis (non nitidis) nerviis secundaris 24-30 (non 18— 24)-
jugis, 6.5—9 (nec 12—13.5) cm latis, petiolis 2.1-5 (non 0.5—1.4) cm longis, pedicellis 0.8—1.4
(non 0.3—0.5) mm longis bracteis florinis pedicellis 6-7 oa 3)-plo longiores statim cognoscitur.
Treelet to 4 m tall. Branchlets terete, 7—9 mm diam., lenticellate, the bark
vertically ridged, densely rufous tomentose. Leaves alternate; blades mem-
branaceous, oblanceolate, 33-49 cm long, 6.5—9 cm wide, apically subacuminate,
basally long-attenuate, dull green above and below, the veins 24—30 pairs,
bullate above, prominently raised below, the tertiary areoles prominently
raised below; glabrate above, moderately rubiginous furfuraceous lepidote
and sparsely pellucid punctate below, the margin entire, decurrent, gradu-
ally tapering to the petiole; petioles marginate, 2.1—-5 cm long, to 0.5 cm
thick, glabrous, bape pulvinate, the pulvinus 1.3—2 cm above petiole
base. Staminate inflorescence: a simple, lax, axillary raceme 6.5—8 cm in bud;
the rachis, bracts and pedicels densely rufous tomentose; floral bracts mem-
branaceous, linear-lanceolate, 34 mm long, 0.3—0.4 mm wide, apically subulate,
the margin entire; pedicels cylindrical, 0.8-1.4 mm long, glabrescent. Staminate
flowers 4-merous; calyx membranaceous, cotyliform, 1.4—1.6 mm long, the
tube 0.2—0.4 mm long, the lobes widely ovate, 1—1.3 mm long, 1.2—1.5
mm wide, apically acuminate, densely and prominently black punctate, sparsely
rufous pubescent, the margin hyaline, flat, erose, epunctate; corolla (in bud)
subrotate, membranaceous, to 1.2—1.5 mm long, the lobes ovate, apically
obtuse, densely and prominently black punctate, glabrous without, sparsely
glandular-granulose within, the margin hyaline, flat, erose, epunctate, gla-
brous; stamens appearing epipetalous, the anthers sessile at the junction of
corolla tube and lobe, deltoid, ca. 0.7 mm long and wide, dehiscent by subapical
pores, the connectives red punctate medially; pistillode, conic, hollow, ca.
0.5 mm long and 0.2 mm diam. Pisti/late inflorescence as in staminate but 3—
6(—7.2) cm long; floral bracts 1-1.6 mm long, 0.2—0.4 mm wide, apically
long-attenuate; fruiting pedicels incrassate, 0.7—0.9 mm long, to 1.5 mm

Pipoty, Cybianthus in Ecuador and Peru 143

Fic. 27. Cybianthus jensonii Pipoly. A. Habit, showing long-attenuate leaf bases. B. Abaxial
leaf surface with prominently defined areoles. C. Portion of staminate inflorescence in bud.
D. Open staminate bud. E. Pedicel, calyx and fruit. A-D, drawn from Gentry et al. 55708.
E, drawn from Croat 19485. Figure drawn by Linda Ellis.

144 Stipa 18(1)

diam. Pistillate flowers as in staminate, calyx 1.4—-1.6 mm long, the tube
0.2-0.4 mm long, the lobes 1—-1.3 mm long, 1.2—1.5 mm wide; corolla,
staminodes and pistil unknown. Fruit depressed-globose, orange, 3-6 mm
long, 6-7 mm diam., smooth, prominently pellucid punctate.

Distribution.—Endemic to lateritic slopes above riparian areas in the
Department of Loreto, Provinces of Alto Amazonas and Maynas, in the northern
Amazon Basin of Peru, 130—210 m.

Ecology and conservation status. —This species occurs in primary lowland
tropical varzea forest margin. It is surely a rare species, given the recent
massive collection effort at the sites during the Florwla of the Biological Re-
serves of Iquitos project (Vasquez 1997), during which the species was not
relocated at the Explorama Inn (Indiana) site. Given its rarity, this species
should be considered threatened.

Etymology.—I dedicate this species to Peter Jenson, President of Explorama
Tours, conservationist and principal promoter of ecotourism in the Peru-
vian Amazon. Much of our knowledge regarding the biology of tropical
ecosystems in the Peruvian Amazon would not have been discovered were
it not for Peter and his associates’ enthusiastic support, generosity and hos-
pitality at the company’s biological reserves (Explorama Inn, Explorama Lodge,
and Explornapo Camp) which now serve as long-term ecological study sites.
Explorama Tours’ properties, with their combination of research, public education
and collaboration with local communities and tourism, serves as the most
successful model for tropical ecotourism known.

Paratypes. PERU. Loreto: Prov. Maynas, Explorama Inn, 2 km W of Indiana on Rio
Amazonas, 03° 30'S, 73° 02' W, 130 m, 12 Feb 1987 (stam. bud), A. cate et al. ae
(AMAZ, MO); Varadero de Mazan from Rfo Amazonas to Rio Napo, 22 Aug 1972 (fr),
Croat 19485 (AMAZ, MO, NY, USM).

Cybianthus jensonii is most closely related to C. grandezii Pipoly, but eas-
ily recognized by its long-attenuate leaf bases, epunctate, pallid abaxial leaf
surfaces, the secondary veins 24—30 pairs, narrower leaves with much longer
petioles, subobsolete pedicels 0.8—1.4 mm long, and longer floral bracts.

39. ar ie a fosteri Pipoly, . nov. (Fig. 28). Type: coe Mapre DE Dios:
ov. Manu, Atalaya, vicinity Hacienda Amazonia, 2—3 km W of village, across Rio
A Macite, 12° 35'S, 71? 12 Ww. forested ridge, 600-900 m, 7 Dec 1983 (stam.
fl), R. Foster & T. Wachter 7254 (HotoryPe: MO; tsorypes: F, NY, USM).
Ob folia | l | b inflorescentia

OL dy 1
| ri | a ]

&

longirace mosa, fl coriaceosque

venezuelano valde affinis sed ab ea ramulis teretes (non angulatis), foliis pseudov erticillatis

(non alternis), laminis denseque manifeste atro- a tatis et omNino prominens (non parcege

plane subter atro cones punctate) pe iolatis canaliculatis (not eat ae ad bases Bpripte
mate

planis)

crassis (ne
(non anne poris separatis (non connec ous) praeclare distat.

Pipoty, Cybianthus in Ecuador and Peru 145

Fic. 28. Cy oe ae Pipoly. A. Habit, showing terete branchlets and pseudovercicillace
phyllotaxy. B. n of staminate inflorescence, showing pustulate corolla and apicallly
rounded es ith separate (not birimose) pores. C. Abaxial leaf surface detail, showing
punctations and sparse puberulence. D. Branchlet apex, showing tomentum. A—D, drawn
from holotype, by Linda Ellis.

146 Sipa 18(1)

Tree to 3m tall. Branchlets terete, (6—)7—9 mm diam., densely ferrugineous
tomentose. Leaves pseudoverticillate; blades chartaceous, oblanceolate, (26—)
28—34 cm long, 5.5—9(—10) cm wide, apically abruptly acuminate, basally
long attenuate, midrib somewhat elevated above, prominently raised be-
low, the secondary veins 1 1—18 pairs, dull green above, pallid below, prominently
black punctate above and below, sparsely rufous puberulent below, the margin
entire, regular, flat; petioles canaliculate, 1.6—2 cm, pulvinate, ferrugineous
tomentose, glabrescent. Staminate inflorescence a lax raceme 8-12 cm long,
moderately rufous lepidote, glabrescent; floral bracts lanceolate, 2—3 mm
long, 0.8—1.1 mm wide, apically acute, densely rufous lepidote; pedicels cylin-
drical, 3.5—5 mm long at anthesis erect in bud, nodding in anthesis, densely
rufous lepidote. Staminate flowers 4-merous, coriaceous, nodding, pale green;
calyx cotyliform, 1.2—1.3 mm long, the tube 0.5—0.6 mm long, the lobes
widely ovate, 0.7—0.8 mm long, |.1—1.2 mm wide, apically rounded, densely
and prominently black punctate, sparsely rufous lepidote, glabrescent, the
margin stramineous, opaque, epunctate, erose-fimbriate, glabrous; corolla
subrotate, 2—2.5 mm long, the tube 0.5—0.6 mm long, glabrous, the lobes
widely ovate, ee mm long, 1.4-1.6 mm wide, apically acute, densely
and prominently black punctate, sparsely rufous lepidote without, glabrescent,
prominently pustulate and densely glandular-granulose throughout within,
the margin stramineous, erose, glandular-granulose; stamens apparently sessile
at junction of corolla lobes and tube, the anthers sessile, very widely ovate,
0.5—0.6 mm long, 0.8—0.9 mm wide, apically rounded, basally truncate,
the pores widely ovate, extending 1/2—3/4 anther length, sey (not confluent),
the connective prominently red punctate; pistillode subglobose, 0.7—0.8
mm long, 0.3—0.4 mm wide, hollow, densely glandular-lepidote. Prsts//ate
inflorescence as in staminate but erect, 3-5 cm long; floral bracts 2-3 mm
long, 0.8-1.1 mm wide; pedicels 2.5—4 mm long in fruit. Pisti//ate flowers
as in staminate but calyx 0.8—1 mm long, the tube 0.2—0.3 mm long, the
lobes 0.5—0.7 mm long, 0.9—1.1 mm wide; corolla, staminodes and pistil
unknown. Fruit globose, 6-8 mm long and diam., red at maturity.

Distribution.—Known only from the type locality, presumably endemic.

Ecology and conservation status. —Cybianthus fosteri is endemic to one area
of the Mant Biosphere Reserve and National Park, one of the largest in
South America. It is a ridgetop species in lowland topical most forest. Given
the extension of the Reserve and the species narrow range, it should not be
considered threatened at this time.

Etymology.—This species is dedicated to Robin Foster, of the Smithsonian
Tropical Research Institute and a research associate of the Field Museum of
Natural History. Robin has served as one of the co-founders of the Rapid
Assessment Protocol, and is an expert on the reproductive biology of Tachigali
(including Sclerolobinm) of the Fabaceae.

Pipoty, Cybianthus in Ecuador and Peru 147

Paratype: PERU. Madre de Dios: Prov. Manu, Acalaya, vicinity Hacienda Amazonia,
2-3 km W of village, across Rfo Alto Madre, 12° 55'S, 71° 12' W, 600-900 m, 7 Dec
1983 (fr), R. Foster & T. Wachter 7242 (BRIT, F, MO, NY, USM)

Cybianthus fosteri is closely related to Cybianthus venezuelanus, but is eas-
ily recognized by its terete branchlets, pseudoverticillate leaves prominently
and densely black punctate above and below, canaliculate petioles, pustu-
late corolla lobes, and rounded anthers with separate (not birimose) pores.
The pustulate corolla lobes are unique within the genus.

40. Cypeot us resinosus Mez in Engl., Pflanzenr. IV. 236(Heft 9):219.
O2. Type: PERU. Loreto: Prov. Mavis near Yurimaguas, without date (fr), E
aoe 2428 (HOLOTYPE: W; ISOTYPE: P).

Tree to 15 m tall. Branchlets 2.5—3.5 mm diam., densely ferrugineous dendroid-
tomentose at first, glabrescent. Leaves pseudoverticillate; blades thinly co-
riaceous, elliptic, (11—)15—21(—26) cm long, (4—)5.5—7(-10.8) cm wide,
apically long-acuminate, the acumen 0.8—1.5(—2) cm long, basally cuneate,
decurrent on the petiole, midrib prominently elevated above and below,
decurrent to petiole base, the secondary veins (9Q—)11—16(—18) pairs, nitid
and glabrous above and below, inconspicuously pellucid-punctate (not vis-
ible when dried), the margin entire, irregular, flat, entire; petioles canaliculate,
1—2(—3) cm long, tapered, densely ferrugineous dendroid-tomentose, gla-
brescent. Staminate inflorescence a lax, simple raceme, 1 1—15 cm long, sparsely
rufous stellate puberulent; peduncle 0.9-1(-1.8) cm long; floral bracts
chartaceous, linear, 1—-1.2 mm long, 0.2-0.3 mm wide, apically subulate,
densely rufous stellate puberulent above and below, the margin entire, early
caducous; pedicels cylindrical, 2.1-1.7 mm long, sparsely puberulent, gla-
brescent. Staminate flowers 4-merous, membranaceous, nodding, greyish-brown;
calyx cotyliform, 0.9-1 mm long, the tube 0.1—0.2 mm long, the lobes
widely ovate, 0.6—0.8 mm long, 0.9-1.2 mm wide, apically acuminate, sparsely
rufous stellate puberulent, densely and prominently black punctate, the margin
hyaline, erose, short glandular-ciliate; corolla subrotate, translucent, 1.6—
1.8 mm long, the tube cylindrical, 0.3—0.5 mm long, the lobes very widely
ovate, 1.2-1.5 mm long, 1.5—1.8 mm wide, flat, apically obtuse, densely
and prominently black punctate and glabrous without, densely glandular-
granulose and pusticulate within, the margin irregular, glandular-granulose,
entire; stamens 0.7—0.9 mm long, the tube inconspicuous, membranaceous,
(.3-0.5 mm long, sessile, the anthers cuadrate, 0.2—0.3 mm long, 0.5—0.6
mm wide, apically truncate, translucent, glabrous, the connective promi-
nently red punctate dorsally; pistillode cylindrical, 0.3—0.5 mm long, 0.2—
0.3 mm diam., hollow, densely punctate, glabrous, the stigma 3-lobed. Pzstz/late
inflorescence as in staminate but (1—)1.5—5 cm long; peduncle (0.3—)0.5—1(—
1.5) cm long; floral bracts 0.6—1 mm long, 0.1—0.2 mm wide; pedicels slightly

148 Sipa 18(1)

obconical, 0.7—1.1 mm long, erect in fruit. Pésti//ate flowers as in staminate

ut 1-1.2 mm long, the tube 0.2—0.3 mm long, the lobes 0.8—0.9 mm
long, 1—1.2 mm wide; corolla 1.3—1.5 mm long, the tube 0.3—0.5 mm long,
the lobes 0.8—1 mm long, 1.1—1.5 mm wide; staminodes as in stamens but
0.4—0.6 mm long, the tube 0.3-0.5 mm long, the antherodes ca. 0.1 mm
long, 0.2—0.3 mm wide; pistil obturbinate, 0.4—0.6 mm long, and in di-
ameter, the style very short, the stigma 3-lobed, the placenta cotyliform,
bearing 2 apically exposed ovules. Frit globose, 5-7 mm diam. at matu-
rity, exocarp black, juicy, edible at maturity. Bisexual inflorescence as in pis-
tillate but a lax, simple raceme, or rarely a poorly formed panicle, S—8(—10)
cm long; peduncle 0.5—1 cm long; floral bracts 1-1.2 mm long, 0.2—0.3
mm wide; pedicels 1.2—1.5 mm long. Bisexwal flowers as in pistillate, but
stamens as in staminate, 0.5—0.8 mm long, the tube 0.2—0.3 mm long, the
anthers ca. 0.3—0.5 mm long, 0.4—0.5 mm wide; pistil as in pistillate, conical,
0.4—0.6 mm long, and in diam. Bisexual fruit globose, 4-6 mm diam. at
maturity, exocarp reddish-black, thin.

Distribution.—Once thought to be endemic to the Iquitos area of Loreto,
Peru, Cybzanthus resinosus is now known (reported for the first time here),
from the Choc6 of Colombia, Amazonian Ecuador, Venezuela, with one disjunct
population in French Guiana (O/deman 3272) growing at 100—200(—-1,300)
m elevation.

Ecology and conservation status. —Cybianthus resinosus is restricted to pri-
mary non-inundated forests on white sand (varillal). While it is locally abundant,
it should be considered threatened due to increasing habitat loss. The Ecuadorean
and Venezuelan populations are unusual because they occur in premontane
pluvial forest and wet forest on lower tepui talus slopes (on sandstone) re-
spectively, each containing numerous lowland elements. It may be expected
in the Rio Cenepa-Rio Santiago Drainage Basins, of Amazonas, Peru, an
area known to show the same environments with numerous pockets of sandstone.

Etymology.—The specific epithet refers to the highly nitid adaxial leaf surface,
giving it a lacquered, resinous appearance.

Specimens examined. ae Valle del Cauca: Bajo Calima Concession, ca. 25
km NW of Buenaventura, 9 km NW of San Isidro intersection on “Canalete,” near gate,

—45° slopes, 03° 59' N, 77° 08' — 50m, 13 Jul 1988 (ster.), D. Faber-Langendoen & J.
Hurtado 1757 (CUVC, MO). VENEZUELA. Territorio Federal Amazonas: Dept ei
base of cliff and forest below slope of Cerro Huachamacari, (3° 39! N, 65° 43' W, |
1,300 m, 5 Mar 1985 (stam. fl), Rk. Liesner 718302 (BRIT, MO, VEN). ECUADOR Na apo
Canton E! Chaco, Rio Granadillo, Campamento de INECEL, “Codo Alto,” 00° 08'S, 77°
28' W, 1,300 m, 13-15 Sep 1990 (fr), W. Palacios 5589 (MO, QCNE). PERU. Hukaues:
Prov. Pachitea, region of Pucallpa, W part of Sirdé Mountains and adjacent lowland, ca. 24
km SE to 26 km ESE of Puerto Inca, from Campamento Sird, 09° 28'S, 74° 47' W, SE to
valley of Rio Negro, 750 m, 29 May 1988 (fr), B. Wal//néfer 14-29588 (BRIT, MO, W,
WU). Loreto: Prov. Maynas, Iquitos, May 1925 (stam. fl), G. Tessmann 5145 (NY); Caseria

ao)

Pipoty, Cybianthus in Ecuador and Peru 149
Mishana, 30 km SW of Iquitos, Callicebus Biological Reserve, 4 km S of Mishana, 19 Aug
1980 (stam. fl), R. Foster 4404 (F-2 sheets, NY); Mishana, 16 Aug. 1978 (pist. Al, fr), J.
Ramirez 17 (AMAZ, MO); Mishana, Rio Saees 03° 50'S, 73° 30' W, 140 m, 16 Aug
1978 (fr), J. Ramirez 132 (AMAZ, MO), Mi along Rio Nanay, 03° 55'S, 73° 35' W,
150 m, 20 Jan 1985 (fr), R. Vasquez GN. ee 6126 (AMAZ, MO, NY); Allpahuayo,
Estacién IAP, 04° 10! S, 73° 30' W, 150 m, 5 Jun 1985 (bud), R. Vasquez ies al. Ooot
(AMAZ, BRIT, MO, NY), 29 May 1990 (fr) R. Vasquez et al. 13764 (AMAZ, MO, USM),
16 Aug 1990 (pist. fl, fr), R. Vasquez & N. Jaramillo 14224 (AMAZ, BRIT, F, aS NY,

EX, US, USM), 4 Dec 1990 (fr), R. Vasquez GN. eee 15237 (AMAZ, BRIT, F, MO,
US, USM), 150-180 m, 29 May 1991 (ster.), R. Vasquez & N. Jaramillo 16681 (AMAZ,
BRIT, aon USM), 150 m, 23 Mar 1992 (ster.), R. Vaaues : cs 17996 (AMAZ, BRIT,
MO, USM); Allpahuayo, ca. 26 km along Iquitos-Nauta Rd., 130 m, 25 Aug 1988 (fr), H.
van der eee (AMAZ, MO); Mishuyacu, near Iquitos, on m, Sep 24—28 1929 (fr),
E. Killip & A. Smith 29873 (E, US), May—Jun 1930 (bisex. fl), G. Klug 1384 (F, NY, US);
Quistococha, 200 m, 27 May 1978 (pist. fl, fr), A. Gentry & N. Jaramillo 22314 (AMAZ,
MO); Altura de Pifia Negra, SW of Iquitos, ca. 3-4 km past Quistococha, 200 m, 19 Nov
1975 (fr); Caserfo de Urcumirafio, Rio Napo, path from settlement to tall forest, 120 m, 8
Oct 1979 (fr), C. Diaz GN. Jaramillo 1486 (MO, NY), Pefia Blanca, on Rio Itaya, 110 m,
19 Sep 1929 (fr), E. Killip & A. Smith 29672 (F, US); Between Yurimaguas and Balsapuerto
(lower Rio Huallaga basin), 135-150 m, 26-31 Aug 1929 (fr) E. Killip & A. Smith 28110
(F, NY, US); Prov. Requena, Dtto. Sapuena, Jenaro Herrera, Rio Ucayali, 04° 55'S, 73°
40! W, 160 m, 16 Aug 1994 (stam. fl), R. Ort/z et al. 101 (AMAZ, BRIT, MO), without
locality, except “in Peruvia subandina, without date (fr), E. Poeppig s.n. (L).

Cybianthus resinosus is most closely related to C. penduliflorus Mart., but is
easily separated from it by the inconspicuously punctate leaves, longer eacS
flat corolla lobes and calyx lobes with acuminate apices and erose, short-
ciliate margins. The adaxial prominently raised midrib decurrent to the petiole
base is unique within the subgenus. The fruit of Cybianthus resinosus is also
smaller, black and has a thick exocarp, and it inhabits serra firme forests on
white sand whereas C. penduliflorus is an igap6 species.

41. Cybianthus fuscus Mart., Flora 259. 1841. Type: BRAZIL. Mato Grosso:
“Prope rivum Cochim in Gat May (pist. fl), P da Silva Manso s.n. (HOLOTYPE: M)
Shrub or small tree to 3 m tall. Branchlets terete, 2—3 mm diam., densely
dendroid and stellate rufous glandular-tomentose, tardily glabrescent. Leaves
pseudoverticillate; blades chartaceous to coriaceous, very narrowly oblan-
ceolate or very narrowly oblong, (13—)16—25(—30) cm long, 2—4(—6) cm
wide, apically long acuminate-attenuate, the attenuated portion 1—2 cm
long, terminating in a minute rounded tip, basally long acuminate-attenu-
ate, the attenuated portion 1.5—2 cm long, giving the petioles appearance
of being longer, fully decurrent on petiole to pulvinus; midrib prominently
elevated above, decurrent to petiole base, the secondary veins 12—25 pairs,
somewhat to deeply impressed, the leaf appearing subbullate to bullate above,
prominently raised and loop-connected below, somewhat nitid and glabrous
above, pallid, rufous Pape and conspicuously black punctate and punc-
tate-lineate below, t rgin entire, flat, glabrous; petiol marginate,

150 Sipa 18(1)

5—10(-12) mm long, with a basal pulvinus. Staminate inflorescence: a lax raceme
(2.5—)5—8(-19) cm long; peduncle 0.8—1.5 cm long; floral bracts chartaceous,
linear, 1.2—1.5 mm long, 0.3 mm wide, apically subulate, densely and promi-
nently rufous papillate; pedicels 3.5—5 mm long, densely papillate . Stami-
nate flowers chartaceous, 4-merous; calyx cotyliform, 0.8—1(—1.8) mm long,
the tube ca. 0.2 mm long, the lobes very broadly ovate or linear-lanceolate,
0.5—0.8(-1.6) mm long, 0.6—0.8 mm wide, apically acute to acuminate to
attenuate, medially thickened, densely and prominently red and black punctate
medially, with a few scattered rufous papillae, the margin scarious, highly
erose, densely glandular-ciliate; corolla subrotate, 2—2.3 mm long, the tube
0.6—0.8 mm long, the lobes suborbicular, unequally divided, 1.4—1.6 mm
long, 1.3—1.7 mm wide, apically broadly rounded, densely and prominently
orange punctate without, densely glandular-granulose throughout within,
the margin often revolute at maturity, irregular, entire, glandular-granulose;
stamens ca. | mm long, the filaments developmentally fused to the corolla
tube for their entire length (the stamens appearing epipetalous), 0.6—0.8
mm long, the anthers very widely ovate, 0.4—0.5 mm long and wide, the
apically acute, basally cordate, apically dehiscent by terminal pores, confluent
at anthesis, ventrally sparsely rufous papillate basally, dorsally densely ru-
fous papillate and sparsely but prominently orange punctate; pistillode absent
or highly reduced, ca. 0.5 mm long. Pésti/late inflorescence as in staminate
but 3—10(-14) cm long; peduncle 6-10 mm long; pedicels 1.5—4 mm long.
Pistillate flowers as in staminate but 0.7—0.9 mm long, the tube ca. 0.1 mm
long, the lobes oblate, 0.6—0.8 mm long, |—1.2 mm wide; corolla as in staminate
but 1.4-1.8 mm long, the tube ca. 0.5 mm long, the lobes ovate to subor-
bicular, 1.1-1.7 mm long, 1.0-1.2 mm wide, the staminodes resembling
stamens but with antherodes 0.3—0.4 mm long and wide; pistil ellipsoid,

—

1—1.2 mm long, 0.6—0.8 mm diam., the stigma capitate, 3—4-lobed, densely
translucent glandular-lepidote, ovules 2—3, immersed in the placenta. Fruit
globose, 5—7 mm long and diam. at maturity, densely and prominently punctate,
with a few persistent translucent lepidote scales.

Distribution.—As here recognized, Cybianthus fuscus occurs ftom the Guianas,
to Venezuela, Colombia, Ecuador, Peru, Bolivia, and their corresponding
frontiers with Brazil. Cybianthus fuscus rims the Amazon Basin, from 100—
200 m elevation. As stated earlier, this is an infrequent distribution, as in
Cybianthus venezuelanus.

Ecology and conservation status. —Cybianthus fuscus occurs in primary forest
on ferra firme, especially on steep slopes, near water courses. It is a locally
infrequent element of the understory but does not appear to be threatened
at this time.

Etymology.—The epithet refers to the color of the tomentum of the branchlets,
pedicels and calyx.

Pipoty, Cybianthus in Ecuador and Peru ia

Representative specimens examined. ECUADOR. Napo: Small area of non- pestis
forest, ca. 60 km upriver from Nuevo Aas 13 Sep 1977 (fr), R. Foster 3618 (F, L
PERU. Loreto: Prov. Alto Amazonas, Capahuari Sur (Petroleum Camp), 02° 51! : o
20' W, 200 m, 25 Mar 1982 (fr), R. et ne 3065 (AMAZ, MO, US); Prov. Maynas,
Rio Yavari, Petropolis, 3 km from Rio Amazonas, 8 Sep 1976 (fr), J. Revilla es (AMAZ,
BRIT, MO); 15 km from roadside along Rd. between UNAP Agricultural Experiment area
and Escuela Forestal Vivero, 9 Feb 1968 (pist. fl), D. Simpson & J. Schunke 647 USM);
Puerto Almendras, 03° 45! S, 73° 25! W, 122 m, 7 Dec 1982 (pist. fl), R. Vasquez GN.
Jaramillo 3511 (AMAZ, MO, NY).

Cybianthus fuscus is a widely defined, infrequent, but widely distributed
species, and is most variable with regard to leaf size and inflorescence stat-
ure. Populations in Ecuador and Peru are almost identical to specimens known
from the Guianas, Bolivia, and Brazil in the northeast portion of Amazonas

state, near the border of Territorio do Roraima. Populations matching the
type have leaves much smaller than the Ecuadorean and Peruvian popula-
tions do, and are more like those of the SE Amazon Basin. While Cuiaba is
located at the northern extreme of the Pantanal Region, it is not entirely
clear where the exact type locality was. If the type locality in what was Cuiaba
Province, was north of the Chapada dos Parecis, then it would be at the
headwaters of the Rio Juruena or Rio Teles Pires, both of which dump into
the Rio Tapajos, then to the Rio Amazonas. If the locality was west of Cuiaba,

toward the Bolivian border, streams there form part of the headwaters of
the Rio Mamasé, a branch of the Rio Madeiras, which empties into the Amazonas
near Manaus. In either case, the type locality would be at the very edge of
the Amazon Basin sensu stricto and it would not be surprising to see the species
in other parts ot the Basin. Therefore, while populations from the type lo-
cality are slightly smaller in stature, the leaves are more chartaceous, and
the inflorescences shorter, there is good evidence to show that they are part
of a large polymorphic ochlospecies complex, of which the populations in
Ecuador and Peru represent a commonly encountered morphotype. The same
pattern of variation seen in this species is seen in many Piperaceae (R. Callejas,
pers. comm.).

Cybianthus fuscus appears to be closely related to C. cuneifolins Mart. (in-
cluding C. indecorus Mez), a vicariant species from SE Brazil. The unique
indument, pedicels obconic in fruit, and striking leaves with very long and
attenuate apices and bases allow for easy recognition of Cybianthus fuscus.
42. Pees cyclopetalus Mez, Bull. Herb. Boissier, Ser. 2, 5:533. 1905.

ZIL. AMAZONAS: near Jurud, Miry, Sep 1903 (stam. fl), z. Ule 5840 (HO-
LOTYPE: ek LECTOTYPE, here designated: HBG; IsoLecrotyres: G, K, M

Shrub to 1.5 m tall. Branchlets terete, 1.5—2.5 mm diam., densely rufous
stellate tomentulose, the comentum appressed. Leaves loosely pseudoverticillate;
blades chartaceous, elliptic to oblanceolate, (9.5—)12.5—16(—20.5) cm long,

152 Sipa 18(1)

3.5—5(—7) cm wide, apically subacuminate to acuminate, basally cuneate,
decurrent throughout petiole length, midrib depressed above, prominently
raised below, decurrent to base of petiole, the secondary veins 10-15 pairs,
dull and glabrous above, dull and sparsely rufous puberulent below, con-
centratec

on

along the midrib and the secondary veins, prominently perpuncticulose
and black punctate-lineate, the margin slightly revolute upon drying, ir-
regular, entire; petioles marginate and canaliculate, (1—)1.5—2(—3) cm long,
tapered, sparsely stellate rufous puberulent, glabrescent. Staminate inflorescence
an erect, simple raceme, (4—)9—1 3 cm long, sparsely rufous stellate puberulent:
peduncle (0.6—)0.8—1(—1.3) cm long; floral bracts coriaceous, linear-lanceolate,
1.3-1.5 mm long, 0.2—0.3 mm wide, apically attentuate to a rounded tip,
glabrous above, densely and minutely rufous stellate comentulose below,
the margin glabrous, entire; pedicels cylindrical, 3.7—5 mm long, sparsely
rufous stellate tomentulose, glabrescent. Staminate flowers 4-merous, coria-
ceous, nodding, green; calyx 0.9-1.1 mm long, the tube 0.2—0.3 mm long,
the lobes linear-lanceolate, 0.7—0.8 mm long, 0.2-0.3 mm wide basally,
apically long-attenuate, densely and prominently red and black punctate,
sparsely rufous stellate puberulent, glabrescent, the margin irregular, erose,
minutely ciliolate; corolla subrotate, 1.5—1.8 mm long, the tube 0.6—0.7
mm long, the lobes very widely ovate, 0.9—1.1 mm long, 1.3—1.7 mm wide,
apically emarginate, densely and prominently red and black punctate, gla-
brous without, densely glandular-granulose througout within, the margin
irregular, entire, flat, densely glandular-granulose; staminal tube inconspicuous,
adnate to corolla throughout, 0.6-0.7 mm long, the filaments short, thick,
0.1—0.2 mm long, glabrous, the anthers widely ovate, 0.5—0.6 mm long,
0.4—0.5 mm wide, apically and basally trunctae, the connective epunctate,
glabrous; pistillode subglobose, 0.3—0.4 mm long, 0.3-0.4 mm diam., hollow,
densely translucent glandular-lepidote, the stigma obsolete. Pistillate inflorescence
as in staminate but 4—6.5 cm long; peduncle 1-1.5 cm long; floral bracts
0.9—1.1 mm long, 0.1—0.2 mm wide; pedicels 1.5—2.5 mm long. Pistillate
flowers as in staminate but calyx 0.8—1 mm long, the tube 0.1—0.2 mm long,
the lobes, 0.7-0.8 mm long, 0.1—0.2 mm wide basally; corolla, staminodes
and pistil unknown. Frvit globose, 0.3—-0.5 mm long and wide, green, exo-
carp thin, red punctate.

Distribution.—Cybianthus cyclopetalus is restricted to the western Amazon
Basin of Brazil and southeastern Peru, to 290 m elevation.

Ecology and conservation status. —This species is retricted to varzea or “tah uampa”
habitats, subject to inundation. Rapid development along the rivers in the
Amazon Basin changes its flow and may effect these populations. There-
fore, it should be considered threatened.

Etymology. —The specific epithet refers to the very widely ovate petals of
the species.

Pipoty, Cybianthus in Ecuador and Peru 153
Representative specimens examined. PERU. Madre de Dios: Prov. Tambopata, Tambopata
Wildlife Reserve, 30 km S of Puerto Maldonado, 12° 15'S, 69° 17! W, 260 m, 9 Nov
1984 (stam. fl), H. Young et al. 146 (MO, US); Tambopata Reserve, at mouth of Rio Orbigny,
12° 50'S, 69° 17' W, 250 m, 6 Mar 1981 (fr), A. Gentry & K. Young 32028 (MO, USM);
Along trail from large laguna at end of Swamp Trail, Explorer’s Inn, near confluence of Rio
Tambopata and Rio La Torre, Reserva Tambopata, 12° 50'S, 69° 20' W, 39 km SW of
ee Maldone 14 Oct 1985 (stam. fl), S. Smith et al, 738 (US), (stam. fl), D. Be// 101
US); Explorer’s Inn, Permanent Plots, Tambopata Reserve, 12° 50'S, 69° 17' W, 290 m,

18 Sep 1994 (stam. fl), R. Vasquez et al. 19132 (AMAZ, BRIT, CUZ, MO).
Cybianthus cyclopetalus is most closely related to C. resinosus Mez, but can
immediately be separated from it by the thinner branchlets, dull, chartaceous
leaves, and the unique linear-lanceolate calyx lobes with long attenuate apices.

43. Cybianthus penduliflorus Mart., Nov. Gen. Sp. Pl. 3:87. 1831 {1829}.
Cybianthus pendulinus A. DC., Trans. Linn. Soc. London, Bot. 17:104. 1834 {orth.
var.}. Cy ce ia A ‘DC. Prodr. 8:117. 1844 forth. var.}. Typr: BRAZIL
AMAZONAS: Prov. Rio Negro, near Ega, 170 m, without date, (stam. fl), C. Moraes
J.2. (HOLOTYPE: a

Cybianthus mac. diate Miq. in Mart., Fl. Bras. 10:292. 1856. Pl. 36. syn. nov. Peckia
macr a Ha (Miq. in Mart.) Kuntze, Rey Gen. Pl. 2:402. 1891. Type: BRAZIL. AMAZONas:

ar Ega, ae ee date, (pist. fl), E. Poeppig 2709 (LecrorypE, here designated: W).

Shrub or small tree to 4 m tall. Branchlets terete, 2.5—3.5 mm diam., stel-
late rufuous tomentose, glabrescent. Leaves alternate; blades chartaceous,
elliptic to oblanceolate, (8—)10.5—19(—28.4) cm long, apically acute, ba-
sally broadly acute, slightly decurrent on the petiole, midrib raised above
and below, the secondary veins (10—)12—15(—23) pairs, nitid above, pallid
below, glabrous, densely black punctate, the margin flat, entire or bearing

a few rough serrulations; petioles semiterete, (0.5—)0.8—1.2(—1.5) cm long,

tapered, glabrous. Staminte inflorescence a lax, simple raceme (8—)10—14(—20)

cm long, densely rufous puberulent; peduncle (0.5—)1—1.2(-1.5) em long;

floral bracts membranaceous, linear-lanceolate, 1—1.2 mm long, 0.1—0.2 mm

wide, apically attenuate, densely rufous puberulent, the margin entire; pedicels

cylindrical, 1-2(—2.5) mm long, densely rufous puberulent, glabrescent.

Staminate flowers A-merous, erect, membranaceous green; calyx cotyliform,

0.8—1.1 mm long, the tube 0.1—0.3 mm long, the lobes ovate, 0.4—0.6 mm

long, 0.6-0.9 mm wide, apically obtuse, densely and prominently black

punctate, the margin crenulate, long glandular-ciliate; corolla subrotate,
1.2-1.5 mm long, the tube 0.3—0.4 mm long, the lobes very widely ovate,
0.9-1.2 mm long, 1.2—1.5 mm wide, apically obtuse to rounded, densely
and prominently black punctate, glabrous without, densely glandular-granulose
throughout within, the margin involute, densely glandular-granulose, en-
tire; stamens 0.7—0.9 mm long, the tube completely adnate to corolla tube,
0.3—0.4 mm long, the anthers ovate, 0.4—0.5 mm long, 0.3—0.4 mm wide,
apically acute to obtuse, basally cordate, the connective prominently black

154 Sipa 18(1)

punctate dorsally; pistillode obsolete. Pestz//ate inflorescence as in staminate
but (2.5—)4—8(—11) cm long; peduncle (0.3—)0.5—1 cm long; floral bracts
1-1.2 mm long, 0.2—0.3 mm wide; pedicels 0.2—-0.5 mm long. Pustzllate
flowers as in staminate but calyx 1.2-1.5 mm long, the cube 0.3-0.4 mm
long, the lobes widely ovate, 0.9-1.2 mm long, 1.2—1.5 mm wide, apically
rounded, corolla and staminodes unknown; pistil conical, |.2—1.3 mm long,
0.9-1.1 mm wide, the stigma 4-lobed, the placenta cotyliform, ovules 4,
naked. Frit 6-10 mm long and in diam., the exocarp thin, densely black
punctate. Bisexual inflorescence: as in staminate but 6-15 cm long; peduncle
(.8—1.2 cm long; floral bracts 1-1.2 mm long, 0.1—0.2 mm wide; pedicels
1—1.2 mm long. Bisexual flowers as in staminate but calyx 0.7—1 mm long,
the tube 0.1—0.2 mm long, the lobes 0.6—0.9 mm long, 0.5—0.8 mm wide,
corolla 1.2—1.5 mm long, the tube 0.3—0.4 mm long, the lobes very widely
ovate, 0.9-1.2 mm long, 1.2—1.5 mm wide, the margin involute, densely
glandular-granulose, entire; stamens 0.7—0.9 mm long, the tube completely
adnate to corolla tube, 0.3—0.4 mm long, the anthers ovate, 0.3—0.4 mm
long, 0.3-0.4 mm wide, apically acute to obtuse, basally cordate, the con-
nective prominently black punctate dorsally; pistil conical, 0.9-1.1 mm
long, 0.7—0.9 mm diam., the style 4-lobed, the placenta cotyliform, ovules
3, naked. Bisexual fruit globose, 4-6 mm long and in diam., the exocarp
thin, pellucid punctate.

Distribution. —C ybianthus penduliflorus is known from Brazil, Peru and Bolivia,
100-200 m. The species is not known from Ecuador, but may be expected
anywhere the auaaae is appropriate within the Ecuadorean Amazon.

Ecology and conservation status. —Cybianthus penduliflorus is endemic to igapo
habitats, aa Wise: flooding. Asa small shrub, it grows on riverbanks
and on hummocks, behind Tyip/aris (Polygonaceae) and other shoreline plants.
At this time, it is not considered threatene

Etymology. 1

- lax habit of the inflorescence,

pendent in the field.

Pe ae. specimens examined. PERU. Loreto: Prov. Maynas, Quistococha, 100
, | Feb 1979 (fr), F Ayala 1623 (AMAZ, MO), Quistococha, 00° 45'S, 73° 15' W, 122
m, 27 Aug 1987 (stam. fl), R. Vasquez & N. Jaramillo 94601 (AMAZ, MO, USM); Lower
Rio Momon, tributary of Rfo Nanay, near Iquitos, 8 Dec 1979 (bud), A. Jones & C. res
sie Susie, CAS, MO), Near Momoncillo, 16 Nov 1976 (stam. fl), J. Revilla 1820 (A
F, MO); Deto. Iquitos, caserfo near Nina Rumi, on Rio — 23 Feb 1976 (fr), J. ai
187 (AMAZ, F, MO, USM), Vicinity of Iquitos, 10 Sep- 12 Oct | alae aus J. Revilla
1442 (AMAZ, BRIT, MO, USM); Morona Cocha, near Iquitos, LOO m, 4 Dec 1962 (fr),
J. Schunke 6268 (AMAZ, F, MO, UCLA, US, USM); Puerto Almendras, ~ ° 48'S, 73° 25!
W, 122 m, 17 Aug 1983 (bud), R. Vasquez & N. Jaramillo 4285 (AMAZ, MO, NY, USM):
Nauta, Quebrada Saragoza, 04° 29'S, 73° 35' W, 150 m, 10 Jan 1988 (fr), R. Vasquez GN.
Jaramillo 10339 (AMAZ, MO, US, USM); Iquitos and vicinity, 11 Oct 1929 (bud), L/.
Williams 3676 (F). BOLIVIA. Santa Cruz: pie ELON ey wane El Refugio, along
Rio Paragtia, SE of the house, 14° 46' 09" S, 61° 11" W, 2 11 Oct 1994 (fl bud),

Pipoty, Cybianthus in Ecuador and Peru Ey

R. Guillén & G. Salvatierra 2290 (BRIT, MO, USZ); Campamento La Toledo, 1,000 m E of
the house, | §, 61° 09' W, 160 m, 21 Oct 1994 (stam. fl), R. Guillén & R. Choré 2459
(BRIT, MO, USZ), 1 km W of camp, on canoe route to Campamento Toledo, 14° 45! 51"
S, 61° 02! 22" W, 30 Jan 1995 (fr), R. Guillén et al. 3114 (BRIT, MO, USZ).

4° 4?!

ACKNOWLEDGMENTS

Support for my studies on the Flora of Peru was provided by a generous
grant from the Andrew W. Mellon Foundation (during my tenure at the
Missouri Botanical Garden), which is gratefully acknowleged. I would also
like to thank the John D. and Catherine T. MacArthur Foundation for their
support of fieldwork in Peru as part of the Documentation of Neotropical
Plant Diversity Project, which I directed from 1990-1994. Thanks are also
due to Michael Dillon (F) and the late Al Gentry (MO) for their collabora-
tion with the Flora of Peru project, to Enrique Forero (COL) and Warren D.
Stevens (MO) whose advice was invaluable. Many thanks to Jon Ricketson
(MO), then project coordinator and now colleague in Myrsinaceae system-
atics and authority on the systematics of Parathesis. For their warm hospi-
tality and collaboration, whether in the field in the herbarium, I express
my sincere gratitude to Rodolfo Vasquez (MO), Camilo Diaz (MO), Nestor
Jaramillo (MO), Cesar Grandez (UNAP), Rosa Ortiz de Gentry (MO), Martin
Timana (TEX), Abundio Sagaéstegui (HUT), José Mostacero (HUT), Joaquina
Alban (USM). My volunteer at MO, Catherine Mayo, assisted in the move-
ment of specimens and recording of determinations. Studies on Cybianthus
subgenera Microconomorpha, Conomorpha, Triadophora and Comomyrsine were
completed during my tenure as a Postdoctoral Fellow in the Department
of Botany, National Museum of Natural History, Smithsonian Institution.
My continued status as a Smithsonian Research Associate has permitted me
to continue studies in the group, particularly in their strong Peruvian and
Colombian collections. I thank Linda Ellis, Peggy Duke, Juan Pinz6én, and
Linny Heagy for the skillfull illustrations of the new taxa described herein.
Iam most grateful to the directors and curators of herbaria cited, especially
F, G, GB, K, MO, NY, S, and US, for their patience and work in arranging
for loans and various transfers of material. Finally, to Jon Ricketson (MO),
and Ted Barkley (KSC) for their painstaking review of the manuscript and
helpful comments, and to Barney Lipscomb for his unselfish, time-consuming,
and meticulous work in editing, I owe my sincerest gratitude.

—

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NUMERICAL LIST OF CYBIANTHUS TAXA

1. Cybianthus pastensis 24. C. pee
2. C. gigantophyllus 25. C. vasquezii

31 Gio eas 26. C. cenepensis

4. C. spichige 27. C. nanayensis

5. C. lepidotus 28. C. marginatus

6. C. laetu 29. C. lineatu

ats peruvianus 30. C. magnus

8. C. comperuvianus 30a. C. magnus subsp. magn
9. C. guyanensis subsp. pseudoicacoreus 30b. C. magnus subsp. asymmetricus
10. C. timanae 31. C. incognitus
11. C. cuatrecasasii 32. C. minutiflorus
12. C. nestorii 33. C. huampamiensis
13. C. spicatus 34. C. granulosus
14. C. fulvopulverulentus subsp. magnoliifolius 35. C. flavovirens
15. C. verticilloides 36. C. venezuelanus
16. C. croatii 37. C. grandezii

17. C. humilis 38. C. jensonii

18. C. sprucei 39. C. fosteri

19. C. simplex 40. C. resinosus
20. C. kayapii 41. C. fuscus
21. C. anthuriophyllus 42. C. cyclopet

2. C. schlimii 43. C. a ieee

LC

. poeppigil

158 Sipa 18(1)
LIST OF EXSICCATAE
Figures in parentheses refer to numbers from the numerical list of taxa.
Collection numbers in boldface type indicate type specimens.

Acosta- rhe. M. 5544 (20); jae M.&D. Soir (7); oo (7); 805 (9); Al bert de Escobar, L.
et al. 3744 (16); Alvarez, A. et al. eos 412 (1); 490 (1); 1330 (1); Ancuash, E. 93 (8); 141 (8);
11 oy 220 (8); 274 (33); 392 en 522 (26); 580 a. 588 3) 731 (334); we (20); 1412
437 (2); André, E. s.n. (22); 1151 (22); 3819 (17): 4551 (1); L. et al. 134 (2 nye royo, L.
K. Keill 164 (8 one L. et al. 510 (9); 674 (8); a und E. oe 20); 10 cae ); Asplund, :
10302 (23); 12497 (23); 18717 (23); Aulestia, C. . 1313 (3); Aulestia, M. & J. hee 925 (9);
Ayala, F. 1623 ( i. 2102 (36); Ayala, F. et al. 2814 ve a :

Balslev, H. 4295 (1); Balslev, H. & Santos Dea 2850 (15); Bang, M. 2048 (36); Barbour, P. 2405
(31); Barbour, P. 2567 (431); Bell, D. 101 (42): Berlin, B. sane 1760 (33); 779 (33); Betancur, J.
et al.4857 (19); Bohlin, a et al. 1493 (18); Boyle, B. 3473 (19); 3599 (19); Boyle, B. & bag Bradford
1878 (19); Boyle, B. et al. 3373 (28); 3450 (30b); eee i etal. 55 (1); ee J. 42095 (1);
Brandbyge, J. eee A. ye 1); Bravo, E. one Gémez 49 (21); Brenes, A. 20530 (22); Buchtein,

753 (8); 1758 (5); Bunting, G. & L. Licht 775 (22); Burger, W. & R. Stolze ee 2):

eng C. et al. 2852 (22); Callejas, R. et al. 2737 (3); Camp, W. E-4809 (28); ee J&L.
(28); Campos, J. et al. 3161 (6); ae de R. 2873 (18); 4746 (22); Castro, J. 27 (7); Ceron, C.
vee 2585 (23); 7184 (17); 7409 - ese & M. Factos 7641 (20); a Ceré6n, C. &
N. Gallo 5063 (9); Chacon, I. say Herrer. 2022), Cogollo, A. & C. Estrada 296 (3); Cogollo, A.
et al. 7529 (3); Collenete, B. 113 (8); Corn — xX & C. I poe 2979 (19); 4339 (19); Croat, T. 7115
(18); 17167 (22); 19485 (38), ae 31); 21194 (8); 37785 (23); 49361 (20); 59009 (16); 72394
(19); 73352 (18); Croat, T. & A. aie ae 8); Caress. 14053 (18); 14138 (18); 14918 (3):
14947 (30a); 15474 (22); 15658 (3); 15743 (18); 16272 (20); 16272 (18); 17312 (22); 17625 (18);
19892 (18); 21721 (15); 21805 (28); 21981 (15); 221 31 (3); 23583 (11); 23734 (3); 23855 (22).

~

Daly, D. et al. 5926 ( Ae Nevers, G. & H. Herrera 6642 (22), Diaz, C. 566 (22); 7265 (20); 1474

(31); 1486 (AO); 1 & J. ake 3711 (1); Diaz, C. & A. Torres 7805 (30a); Diaz, C. et al. 3330

(1); 4448 (1); 6930 (33); 6993 (7); 7007 (2); 7170 (7); 7252 (5 ); 7585 (30b); 7649A (34); Dillon, M.

etal. 121 on en we, ae 9283 (22); Dodson, oe mae 7); Dodson, C. et al. 7597 (19);

a (19 B Domingue, 47 (20); Dressler, R. 60 20); Drew, W. E-35 1 (28); ee
. 10668 oe 0690 » LO803 (6): 10910 (28); 10922 ( i 0931B (28); | 132. 4 (AOb); 1

“ 13124 (13); 13

Encarnacién, F. 864 (7); 26200 (4); Espinal, $. 2109 (22).

Faber- Piet n, D. & J. Hurtado 1757 (40); Ferreyra, R. 1040 (32); Figueroa, A. 875 (15); Folsom,
J. 4544 (18); Fonnegra, R. et al. 3076 (23); re ro, E. & L. coe 7035 (8); 1325 (22); Forero, E.
et al. 7 s eee oe a a mito 1); 20148 (22); 22376 (11); Fosberg, F. R. & Giler, M. 23119

4226 (A); re (27); 4327 (27); 4404 (40); 7817 (32); 8624
(32); 8981 Apes ae ae Fos te 7 & T. Watcher 7242 (39); 7254 (39); Foster, R. & B. d’ Achille
10013 (23); Foster, R. 8962 (8); an (1); 10993 (9); Franco, P. et al. 1059 (22); Franco, P. et al.
3204 (17); 15423 (23); Funck, N. .. Schlim 678 (36).
Garcia, F. & J. Echavarria 259-A (20); Garcia, F. & J. Echavarrfa 259-A (17); Gehriger, W. 471 (28);
Gentry, A. 12419 (30b); BOE): 80096 (4); Gentry, A. & J. Aronson 25044 (2); Gentry, A. &L.
Emmons 38776 (27); Gentry, A. & E. Forero 7317 (18); Gentry, A. & N. Jaramillo 22134 (40); Gen-
try, A. & A. Juncosa 41702 (22); Gentry, A. & S. Mori 13665 (23); 13709 (23); 14023 (23); Gentry,
A. &G. a 26565 (19); Gentry, A. & J. Solomon 44407 (5); 44668 (5); Genery, A. & K. Young

32025 (27); 2 a 2); Gentry, A. et al. 16842 (3); 16867 (3); 21033 (27); 22367 (27); 22859 (33);
22859 (31); 2 1 (31); 27952 (2); 29700 (38); 31479 (27); 35062 (20); 35188 (19); 39040 (7):
40453 (18); Ter 42183 (9); 42247 (20); 51083 (36); 51400 (36); 53167 (3); 5332 oa
$3717 (18); 53881 (11); 53960 (11); 55053 (19); 55708 (38); 72129 (9); 72499 (19); 74630 ( QO);
ae (4); 56531 (4); 53551 (3); Gleason, H. 159 (36); Grandez, C. et al. 1824 (37); 4487 °)

11 (7); 5321 (12); 5370 (12); 5642 (12); Grayum, M. & G. Shatz 3170 (22); Grubb, P. et ae ]

Prpoty, Cybianthus in Ecuador and Peru 159

(6G); 1073 (1); 1633 (20); Guidifo, E. eee Guillén, R. & G. een 2290 (43); Guillén, R. &
R. Choré 2459 (43); Guillén, R. ec al. 3114 (43); Gutierrez, G. 1186 (23).

Haenke, T. 98 (1); Hammel, B. 17150 (19); Harling, G. 25313 (1); 25334 (1); Harling, G. & L
Andersson 17255 (23); 18778 (19); 21594 (1); 23540 (1); 23842 (1); 24097 (23); 9764 (23); Hartman,
R. 2401 (23); ao ae eae 2996 (335); eeu 1200 (28); seve O. 1502 (22);
4660 (22); 5479 (22 Andez, J. & S. Hoyos 483 (22); Holm-Nielsen, L. 6818 (1); Holm-Nielsen,
L. eral. 3965 (1); ee 19295 (18); Hoover, W. 1194 (22); Hoover, W. et on 2456 (19); 2809
(19); 2815 (22); 3358 (19); Huashikat, V. 356 (20); 507 (20); 581 (20); 677 (34); 1221 (34); 1248
(23); 1422 (34); Higabe Idt, A. von & A. Bonpland 1096 (13); Hurtado, F. & D. Neill 235 (3);
Hurtado, F. & A. Alvarado 503 on

Idrobo, J. et al. 9671 (17); Imthurn, E. B/9 (14).

1 tea J. 321 (8); 436 (7); 1351 (33); 3929 (28); 8298 (19); 8501 (9); 8522 (7); 30984 (23); Jaramillo,
J. & E. Grijalva 12988 (3); Jelski, C. von 11 (1); 360 (8); Jones, A. & C. Davidson 9717 (43); Jorgensen,
P. et al. 1278 (28); 1297 (1); Juncosa, A. 1255 (22); 1769 (22).

Kalbreyer, W. 1087 (30a); 1534 (17); Karsten, H. s.n. (22); Kayap, R. 558 (8); 618 (33); 723 (20);
iG a 2); 783 (33); 856 (33); 982 (33); 993 (33); Kegel, H. 2 Sky el 2601 (19); Killeen,
449 (32); 7528 (14); 7530 (14); Killip, E. 39372 (23); 35675 (22); Killip, E. & H. Garcia 33802
a 33886 (1); 33898 (3); Killip, E. & A. Smith 24011 (8); VG EN 26286 (8); 26464
(8); 26548 (8); 26563 (8); 27005 (9); 28110 (40); 29871 (9); 29873 (40); 29906 (27), 33279 (18),
Kirkbride, J. & J. Duke 610 (22); Klug, G. 94 (36); 285 (35); 304 (27); 367 (35); 724 (27); 1384
(40); 1412 (9); Klug, G. 2108 (18); 2565 (9); 2691 (20), ae (2); ee 3981 (32); a S.
8290 (7); Knapp, S. & J. Kress 4282 (22); Knapp, S. et al. 8514 (13); ce ); Kohn 11 (18);
Krukoff, B. 1377 (22); 1388 (22); 7240 (13); 7290 (22); 7663 (20); 7663 (22); 10930 rs 10987
(5); Kujikat, A. 50 (33); 265 (26); 306 (26); 395 (33).

Lawrence, A. 370 (22); ee Lehmann, F. s.n. (17); 599 (28); 2399 (28); 3027 (15); 5143 (1);
6202 (1); 651 (18); Lewis, W. et al. 10180 (18); 10306 (25); 10312 (32); 10340 (21); 10475 (21);
12853 (21); 14051 (21); He oo: Liesner, R. et al. 18302 (40); 15093 (22); Limach, C. 140 (20);
Linden, J. 447 (28); Little, E. 7481 (11); 8532 (3); Lleras, E. et al. P17315 (22); Lowrie, S. et al. 52
22); Lozano, G. 3366 (11); 4133 (11); Lugo, H. 3633 (23); Luceyn, J. & Cotton, E. 11011 (1); 11295
(28); 11414 (6); Luteyn, J. et al. 6647 (28)
Macbride, J. 5677 (8); Maddison, M. & L. Besse 7201 (22); Maddison, M. et al. 3204 oe Malme, G.
ee 3483 (8); Malme, G. 3483 (8); Manso da Silva, P. s.n. (41); Marmillod, R. 4-R-90 (4); R-
7 (4); Martius, C. von s.n. (43) s.n.-1820 (9); s.n.-1826 (13); Mathews, A. s.n. (7): 1561 (6);
Se S. 10942 (7); MEDAN niel, S. & M. Rimachi 31219 (37); gay G. et al. 13212 (1);
13397 (1); oe de 790 (18); 797 (18); 1124 (18); Morawetz, W. & B. Wallnéfer 12-27188
(13); 13-11888 (8); 13—30888 (8); 14-31188 (4); 22-19188 (7); Mori, S. aie (22): Mori, S. & A.
Bolten 7292 (1); see 49 (22); 2919 (22); 3907 (22); 5102 A (22)

Neill, D. 7494 (9); 9602 (10); 9813 (18); Neill, D. & W. Palacios 9518 (30a); 9556 (10); 9615 (7);
Neill, D. et al. 10303 (9).
Ortiz, R., et al. 74 (9); 98 (7); 101 (40).
Palacios, W. 1466 (23); 4471 (18); 5394 (3); 5589 (40); Hea (3); 5950 (17); eer (3); 6176 (3);
ces 6218 (3); 6734 (22); 10680 (18); 11407 (8); Pal s, W. & D. Neill 1584 (21); Palacios,
& G. Tipaz 10569 (28); Palacios, W. et al. 1040 (9); Ten (9): 7761 (7); ns 8346 (10);
. (07 (10): 8483 (22); 8486 (7); 9269 (22); Pavén, J. s.n. (1); s.n. (1); Pearce, R. 250 (28); Pennell,
F. eye (3); Pinkus, A. 181 (13); Pipoly, J., A. Cogollo et Al cee (3); 17182 (3); 21026 (19);
21046 (19); 21051 (19); 21055 (19); 21109 (19); 21131 (19); 21144 (22); 21148 (19); 21149 (22);
21173 (19); 21240 (17); 21241 (20); 21249 (19); 21296 (19); 21326 (19); 21328 (19); 21417 (19);
21469 (19) 21471 (19); 21520 (19); 21524 (19); 21598 (19). Pipoly, J., & A. Cogllo 17322 (1);
17376 (1); 17505 (1); Hee 17534 (1); 17881 (1); Pipoly, J. & J. Murillo 15483 (20); ae Js
et al. 3307 (3); 12110 (7); 12112 (7); 12263 (27); 12383 (20); 12490 (20); 12492 (9); 12497 (2);
12541 (20); 12706 (9); one (2); 13284 (2); 13423 (2); 13426 (2); 13931 (2); 14174 (32); 14 ne

Qo

—~

160 Sipa 18(1)

(7); 14997 ee rae ae (20); 17183 (23); oe : pee 7 17281 (3); 17360 © at
17361 (17); 17406 (22); 42 (3); ree 17979 (3); ee 6 (8); Pitman, N. 688 (

993 (19); 1161 (19); Poeppig, E. s.n. (40); 2428 (40); si oe. 00 (43); Prance, G. et al. ee
(22); P12555 (20); 19075 (8).

Quelal, C. et al. 764 (22).

Ramirez, J. 17 (40); 132 (40); Restrepo, C. 723; Revilla, J. 187 (43); 1302 (41); 1826 (43); Riedel, L.
959 (8); Rimachi, M. 7232 (7); Robert, A. 32 2»? (8); Rodriguez, E. 283 (33); Rodriguez, R. & J. Surubfs
669 (14); Rubio, D. et al. 1335 (20); 1262 (22); 2252 (1); Rudas, A. & A. Prieto 3147 (20); Rueda,
R. & J. Ruiz 597 (9); Rufz, J. & J. Pavén 5/36 (8).

Schlim, L. 686 (22); Schomburgk, ears 885 (13); 1002 (13); Schultes, R. E. & G. Black
8427 (20); Schultze-Rhonhof, H. 2983 (19); Schunke, J. 1049 (8); 3068 (22); 4779 (32); 5308
(36); Slice = 7 ( — 5881 a 6268 (43); 7106 (23); 10883 (35); 10895 - paepatd. J: 323
(22); Silverston P., etal. 2871 (20); Simpson, D. & J. Schunke 647 (41); 784 (9); Smith, D.
aaa ay: As 6782 (10), aa D. & R. Foster 2509 (29): Smith, S. ec al. aa (42); Sneidern,
K. von AG1 2bis (18); 919 (22); 1615 (18); Sodiro, A.; 100/2 (17); 100/12 (19); 100/14 (20); 100/14
(17); Soejarto, D. 3205 (22); 4090 (23); Solomon, J. 3466 (27): 1 5940 (1); 17081 (19); 17091 (19);
ichiger, R. & F. Encarnacién 1224 (4); 1027 (4); Spichiger, R. et al. 1973 Sy: Spruce, R. 1040
(13); paper 5175 (1); 6143 (19); 6144 (18); Stein, B. 3721 (11); Stein, B. & C. Todzia 2292 (6);
Stein, B. & L. Dade 3284 (3); Steyermark, J. 53345 (28); 53550 (3); set (30a); 53642 (1);
53897 (1); oo (11); 54800 (28); 58625 (5); Stork, H. & Horton, X. 10134 (1)
Tate, G. H. H. 741 (5); 927 (5); Terry, M. & R. 1490 (18); Tessman, G. 5145 (40); 3525 (7); 3650
(9); 4493 (33); Timana, M. 1047 (10); 922 (32); Tipaz, G. et al. 260 (22); 1706 (20); 1709 (19); 1741
(30b); 1886 (19); 1950 (22); Todzia, C. & B. Stein 27410 (28); Toro, R. 356 (22); Torres, J. H. 1054
(11); Trail, J. 508 (13); Triana, J. 4 (22); 2562 (3); 2585 (1); 2589 (3); 7594 (22); Tunqui, S. 161
(34); 289 (31); 488 (33); 549 (22); Ule, E. 5160 (23); 5840 (42); 6792 (30b)
Ule, E. 5840 (42); 6792 (30b); 8722 (14); Uribe, L. s.n. (6).
Vargas, I. et al. 2248 (36); Vasquez, R. 8905 (14); Vasquez, R. & J. Criollo 1801 (4); Vasquez, R. &
{. Flores 1824 (37); Vasquez, R. & N. Jaramillo 1993 (25); 2703 (9); 3261 (27); 3511 (41); 4285
(43); 5060 (7); 5075 (27); 5086 (27); 5241 (27); 5867 (7); 6122 (12); 6126 (40); 6137 (4); 6325 (20);
7070 (27); 7570 (7); 7587 (7); 7593 (27); 9461 (43); 10339 (43); 11100 (20); 14092 (20); 14204
(27); 14224 (40); 14465 (27); 14545 (9); 15016 (9); 15237 (40); 16681 (40); Vasquez, R. et al.3065
ve SOL (4); 6551 (40); 6588 (4); 7503 (7); 12108 (2); 13764 (40); 14161 (27); 14335 (7): 17996
40); 18163 (32); 18395 (20); 18520 (32); 19132 (42); 20045 (33)
Wallnéfer, B. 111-1588 (13); 11-16888 (1); 14- 29588 (40); 18-14488 (30b); 112-13688 (13); 12-
17788 (8); Warscewicz, s.n. is Weberbauer, A. 4354 (6); 4668 (7); 4699 (32); 6099 (1); 6121
(28); 7168 (28), Webster, G. 28702 (19); Webster, G. & P. Delprete 27594 (19); Webster, G. & B.
Castro 28769 (19); Webster, G. et al. 27795 Seri ee (19); 28796 (19); Werff, H. van der, et al.
10273 (40); 13074 (11); 12430 (18); 13280 (7); Werff, H. van der & & I. Cabrera 15786 (15); Wiggins,
I. 10484 (28); Williams, LL. 657 (27); 658 (27), 1 o (2), 3676 (43); Wolfe, J. 12346 (32); Wolfe, J
12295 (13); Wurdack, J. 232

‘=

~

Young, H. et al. 146 (42); Young, K. et al. 179 (
Zak, V. 1350 (19); 1545 (19); Zarucchi, J. et al.7201 (3)

NEW FLOWERING PLANTS FROM SOUTHERN
NEW IRELAND, PAPUA NEW GUINEA

W. TAKEUCHI

Botanical Research Institute of Texas
c/o Papua New Guinea Forest Research Institute

P. 0. Box 314, Lae, 411, PAPUA NEW GUINEA
JOHN J. PIPOLY II

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060 U.S.A.
ppipoly @brit.org; clusia@latinmail.com

ABSTRACT

Botanical exploration of the Hans Meyer Range in southern New Ireland revealed the
existence of two previously unknown taxa in the families Rubiaceae and Corsiaceae. The
novelties, Psychotria osiana Takeuchi & Pipoly, (Rubiaceae) and Corsia purpurata vat. wiakabui
Takeuchi & Pipoly (Corsiaceae) are described and illustrated. Taxonomic and ecological
notes are also provided for the new taxa.

ABSTRACT (MELANESIAN TOK PISIN)

Wanpela wok bus long sait bilong botani ibin kamap long ol lain maunten bilong Hans
Meyer, Niu Iland Province, long yia 1994. Dispela wokbus o stadi ibin kamapim tupela
niupela samting. Wanpela em sotpela diwai bilong famili Rubiaisi. Namba tu em wanpela
liklik gras nating bilong famili Korsiasi. Tupela igat nem olsem: Psychotria osiana na Corsia
purpurata vat. wiakabui. Dispela stori i tok klia na soim tu sampela piksa bilong tupela.

INTRODUCTION

Papua New Guinea (PNG) is one of only four countries on earth projected
as retaining most of its original forest cover by the end of this century (Suzuki
1993). Unfortunately, it is also among the least known floristic areas within
the Malesian region (Conn 1994; Johns 1995). In the recent multiagency
Conservation Needs Assessment (CNA), 17 localities of primary conserva-
tion significance were identified for Papuasian forest environments (Beehler
1993). Southern New Ireland was one of the enumerated sites considered
vital to biodiversity preservation and in critical need of current informa-
tion. As a direct consequence of the CNA evaluation, a biological assess-
ment expedition was organized by Conservation International and the Papua
New Guinea Department of Environment and Conservation (DEC) in January—
February 1994. A multidisciplinary compilation of results from that sur-
vey has been prepared as a Rapid Assessment Protocol (RAP) Working Paper

Sipa 18(1): 161-168. 1998

162 SipA 18(1)

by Conservation International (Beehler, in press). Orchid specimens from the
Hans Meyer trip were previously reviewed in a separate specialist account
(Howcroft 1994). Two new taxa discerned during general examination of
the expedition’s botanical gatherings are described here.

RUBIACEAE

Psychotria osiana Takeuchi & Pipoly, sp. nov. me 1). Type: PAPUA NEW
sUINEA. New IRELAND: Hans Meyer Range, pond ne Lake Camp,’ 04° 27.2
S, 152° 56,.489' E, 1,175 m, 29 Jan 1994 “. fr), Vv. ae é J Wiakabu 9650
(HOLOTYPE: LAE; isorypes: A, BRIT, K, L)

Quoad Sti ula elonvata calvptrataqaue stipula ab laminis jivenibus fumpentes, P. veratiandet
} #8 oy Pe ee |

o J. 1
- 9 or.

se ‘
> I
floribus verticellatis (non al is) praeditis, laminis secus s furfuraceo- jepidors Gon

glabris) denique petalis desuper papillosis (non glabris) statim cognoscitur.

Shrub or small tree to 7 m height, 8 cm dbh. Stem straight, isodiametric,
basal swell absent; outer bark brown, smooth, slash and sapwood stramineous
to pale yellow. Branchlets terete, 2-3 mm diam., pale brown, slightly com-
pressed at the summit, laxly pilose at apical neds otherwise glabrescent.
Stipules sheathing, calyptrate, caducous, translucent whitish-green, glabrous,
to 5.5 cm long and 6 mm wide. Leaves opposite, usually conferted; blades
coriaceous, elliptic, 10-13 cm long, 3-4 cm wide, apically acuminate, ba-
sally attenuate, the margin entire, anal nitid and glabrous, abaxially
light green and furfuraceous-t long the midrib, the midrib prominulous
on both sides, more elevated es fae secondary veins 10-13 oblique
lateral pairs, excurrently arcuate and usually closing submarginally, the tertiary

reticulum lax, irregular, bifacially raised; petioles adaxially plane, 10—20
mm long, glabrous. lnflorescence a sessile, terminal, pinnate panicle, ruptur-
ing through the stipule, trichotomous (with three principal branches at base),
the branches with flowers racemose, the primary rachis to 2 cm long, greenish,
somewhat angulate, laxly pilose at nodes, internodes subglabrous or pu-

berulent; floral bracts inconspicuous, not persisting; pedicels 1—3 mm long.
Flowers opposed or in verticels, S-merous, entirely glabrous, apparently iso-
morphic; calyx broadly cupuliform, .5 mm long, 1—2 mm wide, limb den-
ticulate; corolla tubular, 3 mm long and obtusely cylindrical in bud, the
lobes chartaceous, corniculate, elliptic to ovate, 1.5 mm long, 1 mm wide,
spreading or reflexing at anthesis, adaxially papillose; stamens erect, epipetalous;
filaments 1.5 mm long, adnate to the corolline sinuses or slightly below.

the anthers basifixed, oblongoid, 0.7 mm long, 0.2 mm wide, apically and
basally emarginate; ovary umbonate, channelled, bilocular, the style bifid,
slightly exserted, the stigmatic lobes spreading and tuberculate. Drapelet
subglobose, 5-8 mm diam., epicarp glabrous; pyrenes 2, not dorsally ridged,
approximately planoconvex, flat on the commissural face, often with one
pyrene aborted or reduced; endosperm ruminate.

TAKEUCHI AND Pipoty, New taxa from Papua New Guinea 163

1mm

_ 1. Psychotria osiana Takeuchi & Pipoly. A.Branchlet. B. Developing infl
aan stipule. C. Inflorescence; showing main rachis and part of the second Rachie
back. Third axis ne for clarity. D. Flower at anthesis; petals reflexing. E. Partially reas
flower. A-D drawn from holotype.

164 Sipa 18(1)

Distribution and ecology —Known only from mossy montane forest in the
Hans Meyer Range of southern New Ireland. Occurring as a seral element
in gap phase regeneration among mature growth premontane forest stands,
and along forest margins.

Etymology.—It is a great pleasure to dedicate this species to Dr. Osia Gideon,
Deputy Director of the Papua New Guinea Forest Research Institute and a
specialist in Papuasian Rubiaceae and Zingiberaceae, for his relentless ef-
forts to document the rich, largely autochthonous, flora of New Guinea.

Pararype: PAPUA NEW GUINEA. New IRELAND: Hans Meyer Range, pond next to
‘Lake Camp,’ 04° 27.205'S, 152° 56.489' E, 1,175 m, 27 Jan 1994 (fl, fr), W. Takeuchi G
J. Wiakabu 9560 (A, BRIT, K, L, LAE).

Papuasian Psychotria were first revised by Valeton (1927) and the nonclimbing
representatives more recently treated by Sohmer (1988). Infrageneric rela-
tionships are still poorly understood due to the taxonomic difficulty of the
genus and the absence of regionally-based studies. However a number of
informal species groups have been recognized by Sohmer from macroscopic
features such as stipule form and inflorescence structure.

Psychotria osiana is characterized by a contracted trichotomous inflores-
cence (1.e., with 3 rachises branched at the base), flowers opposed or verticelled
along the main axes, leaf blades with abaxially furfuraceous-tomentose midribs,
completely glabrous flowers, and petals adaxially papillose. The corolline
papillae are apparent in fresh or rehydrated material, but less so i” sicco.

A total of 17 species of Papuasian Psychotria have sheathing stipules of
the sort present in Psychotiria osiana. Developing inflorescences and young
leaves emerge by rupturing through the stipule, which subsequently disin-
tegrates or falls away intact. The combination of calyptrate (sheathing) stipules
and a trichotomous inflorescence occurs in only two species: P. leleanoides
Sohmer and P. /orentzii Valeton. Psychotria osiana is clearly related to P. leleanoides,
(a species from New Britain and the Solomon Islands), but is distinguish-
able from it by the leaf indumentum and the entirely glabrous flowers with
papillate corolla. Judging from elevational distributions, Psychotria ostana
is the montane sister species to P. /eleanoides, the latter being primarily a
lowland taxon from elevations below 200 m (Sohmer 1988)

CORSIACEAE
Corsia purpurata L.O. Williams var. wiakabui, Takeuchi & eee vat.
nov. (Fig. 2). Type: PAPUA NEW GUINEA. New IreLanp: Hans Meyer Range,
pond next to ‘Lake Camp,’ 04° 27.205'S, 152° 56.489' E, 1,175 m, 28 Jan 1994 (Al,
fr), W. Takeuchi & J. Wiakabu 9611 (HoLoryPE: LAE, in spirit).

Ad Corsia purpuratam var. purpuratam accedens sed ab ea labello acuminato nec ad basem

extendens praeclare distat.

Terrestrial herb to 13 cm height, sciophytic, erect, all parts glabrous and

TAKEUCHI AND Pipoty, New taxa from Papua New Guinea 165

5mm
Tem

Fig. 2. Corsta purpurata L.O. Williams var. wiakabui Takeuchi & Pipoly var. nov. A. Habit,
dorsal view. B. Habit, lateral view. C. Frontal aspect, petals and lateral sepals. D. Median
sepal. E. Petal. F. Basal callosity, from side. A-F drawn from holotype.

166 Sipa 18(1)

dull reddish-brown. Stems terete, weakly sulcate, the nodes distant. Leaves
4—5, spirally arranged, acroscopic, base sheathing; lamina linear-acuminate,
conduplicate or with margins broadly revolute, chartaceous, 9-17 mm long,
4—6 mm wide, to 6-nerval, sometimes apiculate; venation visible as dark
lines, not raised on either side. F/owers solitary, terminal on the unbranched
stem, bisexual; median sepal flabellate, symmetrical, 16-19 mm long, 20—
23 mm wide, nodding at anthesis, unguiculate, bluntly acuminate at the
apex; the basal callosity linear, narrowly raised by 4 mm above the plane of
the ‘labellum,’ summit corniculate, base abruptly truncate at the claw; median
sepal 1 2-veined, the veins parallelodromous, diverging mainly from the proximal
half of the sepal, once or several times furcate before the margin; petals and
lateral sepals isomorphic, basiscopic and incurved, costate, lanceolate, 4 mm
long, 3 mm wide, acumen 1.5 mm long; stamens 6, in 2 whorls opposite
the perianth segments; filaments resembling the style, 0.6—0.7 mm long;
the anther cells 2, oblongoid, 1.2 mm long; style simple, cylindrical, ca. 1
mm long. Frit not seen

Distribution and ecology.—Known only from the type locality in the Hans
Meyer Range on New Ireland. Corsia purpurata var. wiakabui occurs infre-
quently in mature-growth, primary premontane forest, on its floor, cov-

—

ered by leaf duff. This new variety was encountered as a population flush-
ing after recent rains and thereafter evanescent; all individuals were seen in
comparable maturational states.

Etymology,—The new variety is dedicated to Joseph Wiakabu; botanist,
explorer-collector, and colleague from the Papua New Guinea National
Herbarium.

Paratype: PAPUA NEW GUINEA. New IRELAND: ie pe a ee slopes ee
river valley on the ascent from Mandih lake, 04° 26'S, 59' E, 750 m, 10 Aug 1975,

MAJ.S. Sak etal, 2091 (K, LAE

Corsia was revised by van Royen (1972) in a monograph based on 45 exsiccatae
numbers, an average of less than 2 collections per species. The limited number
of specimens available for study is due to population rarity, the inconspicu-
ous habit of the plants, an ephemeral phenology, and the preference for sheltered
microsites in forest where ambient light conditions favor concealment. Van
Royen (ibid.) commented that encounters with Corsza are often of a fortu-
itous nature; occasioned by understory sun flecks falling on the plants ina
certain way by chance. There is no doubt that the genus is rarely found by
collectors. The intervening years since the initial revision has seen little increase
in tora of specimens, nor is the situation likely to improve dramati-
cally in the future. The difficulty of botanizing purposefully for Corsza dis-
courages the sort of field-based studies necessary for understanding the
morphological variation between populations.

TAKEUCHI AND Pipoty, New taxa from Papua New Guinea 167

According to van Royen’s (1972) conspectus, Corsa is a saprophytic ge-
nus with 25 species distributed through New Guinea, the Bismarck Ar-
chipelago, the Solomon Islands, and Australia. Most of the species are nar-
rowly defined endemics of montane forest habitats, and appear to be restricted
to specific mountains. On the basis of the venation and basal callosity of
the median sepal, the new taxon is related to C. purpurata var. purpurata,
but is distinguishable by the flabellate ‘labellum’ with acuminate apex and
the base truncate rather than extended.

The type locality for var. waikabui is floristically depauperate in com-
parison to similar habitats from mainland New Guinea, and is dispropor-
tionately composed of epiphytic or nonendemic species (Takeuchi and Wiakabu,
in press). Depauperate levels of biological diversity in this montane envi-
ronment was also reported by zoological specialists participating in the New
Ireland survey (Beehler in press, passim). The expedition’s collective find-
ings are consistent with a supposition that the cloudy uplands of southern
New Ireland are of geologically recent origin. In view of the multidisciplinary
results from the recent survey, it is unlikely that future exploration of the
Hans Meyer Range will yield significant numbers of additional novelties.

ACKNOWLEDGMENTS

The Rapid Assessment Protocol survey of New Ireland was funded by
Conservation International and the Biodiversity Support Program. Expe-
dition leader Dr. Bruce Beehler and the Papua New Guinea Department of
Environment and Conservation performed key coordinating roles. The Liz
Claiborne and Art Ortenberg Foundation and the John D. and Catherine
T. MacArthur Foundation have provided ongoing financial support to W.
Takeuchi for studies partially based at the Papua New Guinea National
Herbarium, at the PNG Forest Research Institute, in Lae, in support of his
principal work at the Crater Mountain Wildlife Management Area.

Neville Howcroft prepared the illustration of Corsza purpurata var. wiakabui
and Taik Iwagu the illustration of Psychotria osiana, which we gratefully

acknowledge.

REFERENCES

BrEHLeR, B.M. (ed.). 1993. Papua New Guinea conservation needs assessment, vol. 2. Cor-
porate Press, Inc., Landover, Maryland.

BEEHLER, B. (ed.). In press. Rapid Assessment Program Working Papers, No. 10. A biodiversity
assessment of southern New Ireland, Papua New Guinea. Conservation International.
Washington,

Conn, B.J. 1994. Documentation of the flora of New Guinea. In C.-I Peng and C.H. Chou,
eds. Biodiversity and terrestrial ecosystems. Institute Bot., Acad. Sinica Monogr. 14:123-156.

Howcrort, N.H.S. 1994. Orchid collections from New Ireland. Orchid Res. Bull 3. Papua
New Guinea Forest Research Institute

168 Sipa 18(1)

Jouns, R.J. 1995. Malesia—an introduction. Bot. Mag. 12:52-62.

Royen, P. van. 1972. Sertulum Papuanum 17. Corsiaceae of New Guinea and surrounding
areas. Webbia 27:223-255.

SOHMER, S.H. 1988. The nonclimbing species of the genus Psychotria (Rubiaceae) in New
Guinea and the Bismarck Archipelago. Bishop Mus. Bull. Bort. 1:1—339.

Suzuk1, D. 1993. Time for a change. Allen and Unwin, Sc. Leonards.

TAKEUCHI, W. and J. Wiakasu. In press. A transect-based floristic reconnaissance of south-
ern New Ireland. In B. Beehler, ed. A biodiversity assessment of southern New Ireland,
Papua New Guinea. Rapid Assessment Program Working Papers 10, Conservation In-
ternational, W Washington,

VALETON, T. 1927. Die Rubicdeee von Papuasien. I. Zweiter Teil: Coffeoideae. Bot. Jahrb.

1: ie

THREE NEW SPECIES OF CRATAEGUS (ROSACEAE)
FROM WESTERN NORTH AMERICA:
C. OKENNONII, C. OKRANAGANENSIS
AND C. PHIPPSII

J.B.. PHIPPS

The University of Western Ontario
Department of Plant Sciences
London, Ontario, CANADA NG6A 5B7
R.J. OKENNON
Botanical Research Institute of Texas

509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.

ABSTRACT

ee new species of Crataegus, C. okennonii J.B. Phipps (from series Dowglasianae), C.
okanaganensis J.B. Phipps & O’Kennnon (from Oe newly genre series soll B;
Phipps & O’Kennon), and C. phippsii OK rm
North America. All are striking, widespread taxa with a substantial number of er

Key Worps: Taxonomy, Crataegus, Rosaceae, new species

RESUMEN
res nuevas especies de Crataegus, C. okennonii J.B. Phipps (de la serie Douglasianae), C.
okanaganensis J.B. Phipps & O’Kennnon (de la nueva serie descrita Purpureofructi J.B. Phipps
’Kennon), y C. phippsii O’Kennon (préximo a la serie Mol/es) se describen del oeste a
Norte América. Todos son taxa sorprendentes y de amplia difusi6n con un sustancial numero
de cit
Recent fieldwork with R.J. O’Kennon of Fort Worth, Texas in the American
states west of the Rocky Mountains and in British Columbia for the first
author’s revision of the red-fruited Crataegi (Phipps 1998) of this region
has serendipitously uncovered three new species of hawthorn. The first, Crataegus
okennonii, a new purplish-fruited taxon of ser. Dowglasianae (Rehd. ex Sarg.)
Rehd., striking and obvious in the field, is also usually easily resolved in
the herbarium. Crataegus okanaganensis, rarely collected before our studies,
is also a striking new taxon with brilliant red (late August) to deep purple
(late September) fruit. It is accorded a new series, Pupureofructi, together
with the long-described and long-overlooked C. williamsii Egg]. The third
new species, C. phippsiz, though first collected as far back as 1908, remained
otherwise unnoticed until our fieldwork. It is a very distinct purple-fruited
species related to series Molles.

Sipa 18(1): 169-191. 1998

170 Sipa 18(1)

One of the reasons that these new Crataegus species west of the Rocky
Mountains have been overlooked has been the failure of nearly all collectors
to take note of fruit color (Phipps 1998). Each of the following species is
quite distinct in this respect and we therefore take the opportunity to present
color plates of the three species illustrating this point. Hawthorns are con-
sidered to have mostly black or red fruit. The discovery of these three ex-
tends the number of those which are burgundy to deep purple at some stage
in their development. Another feature emerging from this work are new
centers of diversity for western North American Crataegus appearing in the
northern Okanagan of British Columbia and northwest Montana.

A further reason that these taxa have been overlooked is that in the Pa-
cific Northwest region (to nw Montana) and British Columbia where they
occur there has been a belief that generally, only one red-fruited native spe-
cies, - ‘columbiana Howell, one black-fruited native taxon, C. douglasii Lindl.
(incl. C. swksdorfii (Sarg.) Kruschke) and one introduced red-fruited taxon
(C. monogyna Jacq.) occurred. This belief has persisted (Phipps 1998) in spite
of the widespread presence in this region of C. macracantha Lodd. ex Loud.
In view of this lack of taxonomic inattention, including the disappearance
from the floristic literature of the distinctive C. wi/liamsit, it is perhaps not
surprising that earlier botanists did not recognize potential new taxa in this
region. Consequently, we take this opportunity of reviewing the main western
North American species (west of the Rocky Mountains) ina numerical taxonomic
analysis. This analysis uses one synthetic OTU per taxon, each described
from typical material over 38 morphological characters (Table 1) of the kind
commonly used in Crataegus taxonomy. The eleven native species generally
recognized in the west are all present, together with C. monogyna, a useful
outgroup, and five species from east of the Rocky Mountains (C. mo//is (Torr.
& A. Gray) Scheele, C. submollis Sarg., C. brachyacantha Sarg. & Engelm., C.
schuettei Ashe and C. punctata Jacq.) to give structure and scale. Also the
three new taxa described in this paper are included. A dendrogram was prepared
by NT-SYS using the options taxonomic distance and unweighted pair-group
arithmetic averaging. The results (Fig. 1) place C. monogyna and C. punctata
outside all the other species. Among results of general interest, varieties
C. chrysocarpa Ashe var. chrysocarpa and C. ¢. var. piperi (Britton) Kruschke)
uve at about the 0.6 level while distinct species cluster at the 0.8 or greater
level, e.g. C. schuettei and C. wootoniana Egg. (series Tenutfoliae), C. suksdorfii
and C. douglasii (series Douglasianae) and C. erythropoda Ashe and C. rivularis
Nutt. (series Cerrones), Other pertinent results are discussed under the new
taxa described in this paper.

a

TAXON DESCRIPTIONS

1. Crataegus okennonii J.B. Phipps, sp. nov. (Fig. 2). Typr. CANADA.

Puipps AND O’KENNOoN, New species of Crataegus from Western North America tly

Taste 1. The 38 characters scored for the numerical taxonomic analysis.

Plant, general ee lasakaas

ik t habit 22. Flower: r

2. Bark type 23. Pedicel: eee

3. Branch arrangement 24. Anthesis time
Thorns and Twigs Flower

4. Thorns: indeterminate present? 25 ae

5. Thorns: ae 26. Calyx lobes: margins

6. Thorns: curvature 27. Stamen: no.

ie on ae browns 28. Anther: color

8. Thorns, color: gray 29. Style: number

9. Twigs, | yr old, color; browns

10. Twigs, 1 yr old, color; gray Fruit

30. Fruit: color, month before full ripeness

Leaf 31. Fruit: color at full ripeness
11. Lamina: length 32. Fruit: shape, 1

12. Lamina: length/breadth 33. Fruit: shape, 2

13. Lamina: position of widest part 34. Fruit: pubescence

14. Lamina: venation, number 35. Fruit: calyx orientation
15. Lamina: veins to sinuses 36. Fruit: length (height)
16. Lamina: lobe number 37. Pyrenes: lateral faces
17. Lamina: lobe shape

18. Lamina: max. sinus depth (LII) Autumnal foliage

19. Lamina: abaxial pubescence, young 38. Color

20. Petiole: glands

21. Leaf oe glands

BritisH COLUMBIA: pees Valley, E side of 97A 0.5 mi N of intersection with
97, scrubby slopes with long cleared area, 50°21'N, 119°15'W, 19 Aug shee pee
J.B. & O’Kennon, RJ 6998 (HOLOTYPE: UWO; IsoryPEs: BRIT, CAN, TRT, US).

Frutices grandes vel arbores parvae, plerumque trunco uno, ad 10 m alti, spinosi; ra-
muli unius anni nitentes atro-castanei, ramuli veteriores brunneo-cinerei; spinae plerumque
< 2.0 cm longae, crassae, plus minusve recurvatae, in juventute eenaecee itenter. Folia

ecidua; petioli plerumque 1—2 cm longi; laminae 4—6 cm longae, ellipticae vel late-ellipticae,
pene a aaere anguste ovatae vel rhombeae, paribus 4 venarum secondariarum longiorum
ve ybatae vel vix-lobatae vel vadoultee 2—3(-4) lobatae per latus, eae
cea + coriaceae, in maturitate enue appresso- pecs supra, subter + glabrae

venis exceptis, colore autumnali ¢ el pallide carmineum. eee

qua nde

ioe

|

—20 floratae, bracteolis parvis caducis glandulo-marginatis, glabrae vel raro pubescentes.
Flores 15—20 mm diam.; hypanthium glabru m extrinsecus; lobi calycis 3 mm longi, ob-
scure glanduloso- eae petala + circularia; stamina 10-12, antheris aa roseis; carpelli
et styla 3-4. Fructus gi desupialss orcas ca 10 mm diam., triste-purpurei vel nigri
subinde in maturitate, in mense vinosi vel ecicece castanei, reliquiis calycis
parvis; pyrenae 3—4, vadositer sulcatae dorsaliter, lateril m foveis vel interdum planis.

Large bushes or small trees, usually single-trunked, up to 10 m tall, thorny;
one year old twigs deep shining mahogany, older branches gray/brown; thorns
usually < 2.0 cm long, stout, slightly recurved, shiny brown when young.
Leaves deciduous; petioles usually 1-2 cm long; blades elliptic to broad-

Sipa 18(1)

New Crataegus species of western North America

aoustest UPGMA of taxonomic distance
OKENNONII 19 taxa
38 characters
suksdorfii

__J

mollis
submollis
chr.chry

chr piperi %,
macracantha
PHIPPSI!
brachyacantha
saligna
punctata

ee ed y y 7 ¥ T Tr T 7 T T Pe)
0.80 1.20 1.60 2.00

oo
Fic. 1. Dendrogram of western North American Crataegus taxa showing relationships of
species described in this paper. Newly described taxa in capitals. See text for more infor-
mation.
elliptic, sometimes somewhat narrow ovate or rhombic, 4-6 cm long, with
four pairs of longer secondary veins, barely lobed to shallowly 2—3(-4) lobed,
the margins also serrate, somewhat coriaceous, at maturity finely appressed-
pubescent above, below + glabrous except on the veins; autumnal color plum
to crimson or light crimson. Inflorescences 12—20 flowered, with small caducous
gland-margined bracteoles, glabrous to rarely shortly pubescent. Flowers
15-20 mm diam.; the hypanthium externally glabrous; calyx lobes 3 mm
long, obscurely gland-margined; petals + circular; stamens 10-12, anthers
pale pink; carpels and styles 3—4. Fruit + ampulliform-orbicular, ca. 10 mm
diam., dull-purple or occasionally black at full maturity, red-burgundy to
chestnut or reddish-brown in August, calyx remnants small; nutlets 3-4,
shallowly grooved dorsally, sides variably eroded or sometimes plane.
Distribution.—Crataegus okennonit is fairly common in southern interior
British Columbia and the eastern half of northern Washington, ranging to
Idaho and nw Montana (Fig. 3), usually along streams or other sources of
water. It is a tall distinctive plant, usually with a straight trunk, easily rec-
ognized in the field both in flower and in fruit. This new species is most
closely related to C. douglasii, sens. str., from which it differs in fruit shape

PHIpPs AND O’KENNON, New species of Crataegus from Western North America 173

Fic. 2. Line drawing of C. okennonii J.B. Phipps. Inflorescence, flowers and parts from Phipps
6959 (UWO); infructescence, fruit and parts from Phipps 6991, 6995 and O’Kennon (UWO).
Scale bars = 1 cm.

174 Stipa 18(1)

\
oY ‘
) -1239) 1 120 “17 | & Jfyii4

Fic. 3. Distribution map of C. okennoniz J.B. Phipps.

and color, flower size, color of mature leaves (often yellowish-green), habit,
thorn length, leaf shape and autumnal color of foliage.

Crataegus okennoni1 1s one of asmall group of ‘black-fruited’ (fruit + black
at full maturity) species comprising series Dowglasianae (Rehd. ex Sarg.) Rehd.
which I restrict to C. douglasii Lind|., its immediate relative C. suksdorfii
and the new species described here. A greatly widened concept of C. douglasii
which includes C. sa/igna Greene and C. rivularis is advocated by Welsh
(1982) and Holmgren (1997) but in my opinion this cannot stand up due
to many significant differences in leaf-shape, venation type, bark and thorn
type. Moreover intermediates do not exist between sa/igna-rivilaris and douglasii-
okennonii-suksdorfii. Difficulties for the broad concept are also emphasized
by the fact that C. saligna (called C. douglasii Lindl. var. duchesnensis Welsh
by Holmgren) has 20 stamens and cream anthers, contrary to the Intermountain
Flora. Iam (JBP, 199X) therefore placing C. saligna and C. rivilaris ina
different grouping.

Crataegus okennonii is a Clearly defined segregate of C. douglasii differing
by a considerable number of correlated characters of which thorn length
(the shortest in series Dowglasianae) and flower size (largest in the series) are
the most unambiguous. The fo lowing couplet separates C. okennonii from
the restricted interpretation of typical C. dovg/asii as found in western Canada
and the adjacent United States.

Puipps AND O’KENNoN, New species of Crataegus from Western North America L735

1. Usually tree-like with a straight trunk, S—10 m tall; young twigs deep glossy
purple-brown; thorns usually 1.5—2 cm long; flowers ca. 15-20 mm diam.;
fruit crimson lake or chestnut-colored in late August, becoming deep plum-
black at maturity, not bloomy, broadly ampulliform; autumnal foliage usu-
ally crimson, plum or pale shades of same; styles and nutlets usually 3.

C. okennonii

1. Usually bushy, though often large, not conspicuously single or straight-trunked,

3-8 m tall; thorns often longer, to 2.5(—3) cm long; young twigs mid to
deep glossy brown, only occasionally as dark as C. okennonii; flowers often
12-15 mm in diam.; fruit variably colored in late August though never chestnut,

shape ellipsoid or suborbicular; autumnal foliage color often bronzy, not
consistently + crimson or paler; styles and nutlets 3—5. C. douglasii

The differences between C. okennonii and a somewhat restricted C. douglasii
are not by any means large but recognition at the specific level is also sup-
ported by the numerical analysis (Fig. 1), the lack of intermediates among
good quality specimens both in the field and herbarium and the consequent
immediacy of recognition. A more elaborate comparison with C. douglasii
in which detailed leaf shape comparisons will be used may be expected consequent
on the publication of R. Dotterer and T.A. Dickinson’s detailed biosystem-
atic studies of C. okennonii and C. douglasiz.

It is possible that C. okennonii is the same as C. douglasii f. badia Sarg.
(1907), a taxon in which the fruit is described as “chestnut”, certainly not
a common color in ripening fruit of section Douglasii. However, some of
the unripe fruit of C. okennonii seen by us certainly was the color of reddish-
brown, highly polished, shoe leather. Piper, the original collector of forma
badia, variously described the fruit as brown, dark shining brown, chest-
nut, etc. Forma badia is a taxon systematically disregarded in the floristic
literature. Both color changes during the ripening of western North American
Crataegus fruit and the final color of the autumnal foliage have been poorly
understood (Phipps 1997) as taxonomic characters so we wish to record them
here. Crataegus douglasii fruit is shown in Figures 4d and 4e. The latter,
photographed in August, illustrates the black, pendant, ellipsoid fruit while
the other is another British Columbia specimen showing plenty of bloom.
Figures 4a—4c illustrate C. okennonii, plate 4a showing the late August “chestnut”
color while Figure 4b is a later version of the same color. Figure 4c shows
the final, near ripe color, in late September, close to that of C. douglasii. The
foliage is frequently a distinctive bright pale crimson color in late Septem-
ber as is shown in Figure 4f. Variation in leaf form of C. okennonii tends to
fall into two classes, both illustrated in Figure 2, there being a broader,
somewhat rhombo-deltoid, distinctly lobed form (Fig. 4a) and a more el-
liptic or narrow-ovate, shallowly lobed shape (Fig. 4b). Crataegus okennonti
also bears conspicuous, expanding, caducous, coral-red bud scales.

I J.B.P.) am pleased to name this distinctive and handsome new species

176 Sipa 18(1)

aR

Fic. 4. Crataegus okennonii compared with C. douglasit: a) C. okennonii showing remarkable
chestnut-coloured fruit, Kelowna, BC, 18 — 995; a C. okennonit, form with pale foli-
age, note ampulliform fruit, northern Okanagan, BC, 20 Aug 1995; c) C. okennonii with
some fruit close to the final purple-black color, pels Okanagan, BC, 1¢ oe d)
», douglasit showing full ripe color and a high bloom, northern Olanasan, BC, 19 Aug
He e) full ripe color of C. douglasii with ellipsoidal fruit, northern Okanagan, . 19
Aug 1995; f) C. okennonti showing typical cerise ne of full ripe foliage; JBP i in Parentoutie:
Palmer Lake, Washington State, 28 Sep 1993.

after Robert (Bob) J. O’Kennon whose companionship and critical abilities

as a field botanist helped to make this discovery possible. It is intriguing

that so widely distributed a taxon, of which we are able to cite 36 speci-
mens, has remained undetected for so long.

Additional specimens examined: CANADA. BRITISH COLUMBIA: Northern Okanagan,

Soyer: see tees Powerhouse Rd., E side, near Stardel Drive, alt. ca. 1350 ft,

50°27'N, OW, ~ ,4.5 m tall, + erect; fruit deep plum/chestnut, ellipsoid-turbi-

, 19 eee oe SB. sae 164 (UBC, UWO),; Okanagan Valley, Oyama, ca. 0.5 mi

- of E end of spit, alt. ca. 1320 ft, SO°O7'N, 119°21'W, hedgeline on clay soil, bush 5 m

Puipps AND O’KENNON, New species of Crataegus from Western North America 177

tall, fruit plum-purple, turbinate, 17 Aug 1995, ].B. Phipps 7158 (UWO); Castlegar, Selkirk
College grounds, towards bottom of slope beyond beehives, scrubby area, 49°19'N, 117°38'W,
alt 950 fe, tree tall; stamens 5, anthers pale pink, buds forced til 10 May, then 10
stamens seen, 06 May 1994, J.B. — 6928 (BRIT, CAN, DAO, TRT, UBC, UWO, US);
Okanagan Valley, Westside Rd., ca. 9 mi W of jct. with 97 and + due opposite Vernon,
wooded edges on west of road, 50°l S'N, 119°27'W, ca. 1200 ft, tree, 5 m tall, fruit small-
ee pas red- ci tee 19 Aug 1994, Phipps, J.B. & O’Kennon, R.J. 6990 (CAN, TRT,
C, UWO); Northern Okanagan, Hwy 97, E side, between Westside Rd. and Silver Creek
ee alt.1800 a equals JBP 6969, cad 17 m tree, 50°23'N, 119°17'W, convex light
yellow-green foliage, fruit light red, turning to burgundy, roundish, fatter at base, glossy,
19 Aug 1994, Phipps, J.B. & O’Kennon, i 6995 (BRIT, UWO); Salmon Arm, large haw-
thorn pasture just N of railroad and W of central business section, alt. ca. 1900 ft, 50°43'N,
9°16'W; bush, 4.5 m tall, fruic burgundy, locally frequent, (equals JBP 6959); 20 Aug
1994, Phipps, : B. & O’Rennon, RJ. 7027 (CAN, TRT, UBC, UWO); Okanagan Valley,
Westside Rd., ca. 9 mi W of jet. with 97 and + due opposite Vernon on E side of road,
fenceline 50°15! N. 119°27'W; 7 m multitrunked tree, fruit Sees alt. ca. 1200 ft,19
— fee Phipps, J.B. & O’Kennon, RJ. 6991 (BRIT, CAN, TRT, UWO); Okanagan Val-
ley, of Vernon at jet. 97/97A, ee hillside E side of road, a Sep sate J.B. Phipps
ae Pr , ’Kennon 6821 (BRIT, UWO). Pass Creek Rd., N of Castlegar, 1.3 mi along, mesic
grassy slopes, alt. ca. 1800 ft, ete L9O'N, 117°40'W, bush, 4 m tall, fruit ech bacon,
(somewhat ‘chestnut’-ROK), 23 Aug 1994, Phipps, J.B. & O’Kennon, R_J, 7056 (TRT, UWO);
Castlegar, Selkirk ae path behind beehives towards river edge, open woodland-scrub,
alt. 950 ft, 49°19'N, 117°38'W, cree, 20 ft tall, reddish- Ste foliage, dark purple, +
round ee 20 stamens, a Sep 1993, Phipps, J.B. & O’Kennon, RJ. 6824 (UWO); Slocan
V of Slocan, above swamp along small road off Hwy. 6 & just N of Perry Siding
across river, 49°40'N, 117°30'W, hedges and brushy slopes, alt. 1775 ft, bush 6 m tall,
fruit red-burgundy, 22 Aug 1994, Phipps, J.B. & O’Kennon, R.J. 7047 (UWO); NNW of
Vernon on W side Rd about 2.5—3 km SW of intersection of 97 & 97A, alt. 450 ft, 50°20'N,
119°20'W, back of field on west side of road, bush 5 m tall, 24 Sep 1993, Phipps, J.B. G
O’Kennon, R.J. 6785 (UWO); Okanagan Valley, Hwy. #97, ca. 2 mi S of Silver Creek Rd.
exit, (200 m S of ‘road narrows’ sign), (100 m S of JBP 6968), 50°23'N, 119°17'W, alt
L800 ft, in cut-over forest, young tree 7 m tall, 11 May 1994, J.B. Phipps 6969 (BRIT,
TRT, UWO); Okanagan Valley, ca. 8 km NNE of center of Vernon, Pleasant Valley Rd.,
behind Vowle’s residence, hedge at S boundary of Burke’s PYO, 50°18'N, 119°14'W, alc.
380 m, dense, broad, mature hedge, dominant species, 3.7 m bush, 10 mae anthers, thorns
long for species, 2 May 1994, J.B. Phipps 6879 (UWO). Salmon Arm, large hawthorn p
ture, just N of railroad and W of central business center; 50°43'N, 119°16'W, alc. en
ft, bush, 5 m tall, 10 stamens, pink anthers, 10 May 1994, J.B. Phipps 6959 (TRT, UBC,
UWO, V); Okanagan Valley, Newport Beach campsite on W side Rd., approx. due NW of
Vernon, end of parking lot, S end of camp, 50°20'N, 119°22'W, below trees, alt. 1140 ft,
tree 5 m tall, erect habit, burgundy fruit, short thorns, 19 Aug 1994, Phipps, J.B. & O’Kennon,
R.J. 6988 (BRIT, DAO, UBC, UWO, V); NNE of Vernon, S of jct. of 97 and 97A just N
of Baker Hogg Rd., bottom of hill, 50°21'N, 119°15'W, alt. 350 m, 4 m bush, purplish
fruit, 23 Sep 1993, Phipps, J.B. & O’Kennon, RJ. 6821 (UWO); Okanagan Valley, E side of
97, half mi N of intersection with 97; scrubby slopes with long cleared area, 50°21'N,
119°15'W, most abundant species here, bush 6 m tall, red fruit; 19 Aug 1994, Phipps, J.B.
& O’Kennon, R.J. 6999 (UWO); Castlegar, grassy floodplain ENE of Selkirk College, dense
hawthorn thickets on banks of old oxbow, alt. 950 m, 49°19'N,117°38'W, tree, 9 m tall,
foliage elliptical, slightly lobed, purple, fruit globose, dark purple, ?=20 stamens, 27 Sep
1993, Phipps, J.B. & O’Kennon, RJ. 6827 (CAN, ID, MONTU, TRT, UBC, UWO, US);

178 Sipa 18(1)

Northern ay ae ca. | miS of Enderby, gravel driveway to NE starting at Indian Cem-
etery, alt. 1300 fr, 50°31'N, 119°08'W, bush, 4 m tall, fruit burgundy, only one here, 20
Aug 1994, Phin. JBLEO aa RJ. 7006 (UWO).

U.S.A. IDA Idaho Co.: US 12, ca. 10 road mi E of Kooskia, bank at back of field,
N side of road, te arge io 46°07" N, 115°50'W, alt. 1550 ft, tree, pe m call
fruit es 26 Aug 1994, Phipps, J.B. & O’Kennon, RJ. 7089 (UWO). M TANA.

Flathead Co.: Few mi E of Columbia Falls, at dead end of Mt.Creek Rd. (off nae Rd.),
on Calle eae Shoal 330 sign, 48°22'N, 114°08'W, alc.3000 ft, bush, 2 m tall, 08
May 1994, J.B. Phipps 6935 (UWO). Lake Co.: Route 211, ca. 4 mi S of Ronan, nr. MP
44, at farm entrance, driveway through pasture, near scenic turnout, alt. 3030 ft, 47°28'N,
114°06'W, grove of hawthorns on bank S side of driveway, 20 ft apple-like tree, reddish
foliage, black orbicular fruit, 28 Sep 1993, Phipps, J.B. & O’Kennon, R.J. 6841 (UWO);
Rte. 93, ca. 4 mi S of Ronan, near Milepost 44, near scenic turnout, one of group on § side
of farm drive, 47°28'N, 114°06'W, alc. 3030 ft, tree 5 m tall, quite large buds forced ‘til
11 May, stamens 10, anthers pink, (equals JBP 6841), 08 May 1994, J.B. Phipps 6940 (BRIT,
MONTHU, TRT, UWO, US). Sanders Co.: Route 200, 30 mi W of 2 on N side of road
above Flathead R., in group of hawthorns, alt. 2840 fr, 47°19'N, 114°43'W tree, 11 m
tall, + orbicular, bl lack fruit, 29 Sep 1993, Phipps, J.B. & O’Kennon, : 685 58 (BRIT, CAN,

AO, O, V); Route 200, 50.6 km W of jet. with US 93, group of hawthorns
above Hinthead R., 47°19'N, 114°43'W, alt. 2840 ft, tree 9 m tall, stamens 10, anthers
ee e ar IPE & RO’K 6955), 09 May 1994, J.B. Phipps 6955 (UWO); Route 200, 25
3 on N side of road above Flathead R., in long hawthorn hedge, alt. 2845 fc,

47° LNs 114938! W, bush 6 m tall with spheroidal a fruit, 29 Sep 1993, te
J.B. & O’Kennon, RJ. 6857 (UWO); Route 200, exactly 42.6 km W of jet. with US 93,
long hawthorn hedge, above Flathead River, 47°19! N. | . 4! ses alt. 2840 fr, aaa JBP
& RO’K 6857), 09 May 1994, J.B. Phipps 6953 (BRIT, MONTU, UWO). WASH-
INGTON. Ferry Co.: Hwy. 21, ca. 3 km SW of Danville, ies ca. 500 m, 48°59'N, 118°31'W,
marshy roadside with trees and sc ib , tree, 7 m tall, fruit burgundy, (C. douglasiz very common
here), 23 Aug 1995, J.B. Phipps 7174 (UWO). Okanogan Co.: Chopaka Rd., 3 km di-

rectly NW of Palmer Lake, open thicket, low rocky slopes, edge of floodplain, alt. 340 m,
48°56'N, 119°41'W, tree, 5.5 m tall, fruic plum-red to burgundy, 28 Aug 1994, Phipps,
J.B. & O'Kennon, RJ. 7105 (UBC, UWO, WS); Okanogan Valley, W side road about 7 km
N of Tonasket, huge hawthorn stand along riverside, ca. 975 ft, 48°44'N, 119°25'W; bush,
6 m tall, crimson foliage, no fruit, 3 a 1993, Phipps, J.B. & O'’Rennon, RJ. 0865 (UWO);
aa Valley, ca. 5 mi S of Loomis, in depression near roadside, alt. 450 m, 48°45'N,

°37'W, bush, 4 m tall, black fruie, 28 Aug 1994, Phipps, J.B. & O’Kennon, RJ. 7109
ao: ca. 6 km NE of Omak, dense hedges in valley-bottom among farm lands, alt. 250

m, 48°27'N, 119°28'W, dense bush, 4 m ecall, fruit red, changing from orange, 28 Aug

1994, — B. & O’Kennon, RJ. 7100 (UWO). Whitman Co.: US 195 along Spring
Flat Creek few mi S of Colfax at MP 35.3, alt. ca. 775 m, 46°51'N, 117°21'W, tree, 6m
tall, a plum-red (some approaching chestnut), group of same, 28 Aug 1994, —
J.B

—

. & O'Kennon, R.J. 7099 (UWO), 66 Staley Rd., 1 km NE of Chambers, alt. 775
roadside, young tree in front yard of Harold Bough, 46°39'N, 117°11'W, bush, 4 m vi
fruit shrivelled, 27 Aug 1994, Phipps, J.B. & O’Kennon, R.J. 7096 (UWO).

2. Crataegus okanaganensis J.B. Phipps & O’Kennon, sp. nov. (Fig. 5).
Type: CANADA. BritisH ieee Rough grass with hawthorns on Hwy. 97 near
entrance to Kelowna airport, alt. 1800 ft, equals JBP 6907, bush, 4 m tall, fruit,
slightly dull and deep red, + eee 17 Aug ae oo J.B. & O’Kennon, RJ.
6974 (HOLOTYPE: UWO; Isoryprs: CAN, TRT, UBC, US).

Puiprs AND O’KENNOoN, New species of Crataegus from Western North America 179

Y
Af

i
1
Y

Fic. 5. Line drawing of C. okanaganensis J.B. Phipps and O’Kennon. Inflorescence and flower
parts from Phipps 6929 (UWO)); infructescence and fruit parts from Phipps & O’Kennon 7002
and 6975 (UWO); and sterile shoot from Phipps 6894a (UWO). Scale bars = 1 cm.

180 Sipa 18(1)

Frutex, 3—6 maltus, ramuli unius anni mediocriter brunnei vel atrobrunnei, in juventute
sparsim p a Peace spinae validae, 2-4 cm longae, in juventute nitenter atrobrunneae,
recurvatae. Folia decidua, petiolata; petioli 1-2 cm longi, sparsim | ae eglandulares
vel cum glandulis nigribus in juventute (Creston); laminae 3.5—6 cm longae, ovatae vel
ovato-rhombeae, aliquando late ellipticae vel anguste ovatae, lobis 3-4 + acutis, marginibus
4—5 per latus, aliquantum impressis supra, in juventute dense

A

jon

subtilicer dentatis, venis
appresso- pilosis, pilis attritis in maturitate, subter pubescentes praecipue in venis, coriaceae,
super nitentes praesertim in juventute, crescentes rubrae, aeneae in autumno. Inflorescentia
panicula convexa, 10—20 florata; ramuli varie villosi, glandulo-maculati, bracteolis linearibus
caducis glandulo-marginatis. Flores 12—15 mm diam., conspicue cupuliformes in juventute;
hypanthium externe villosum solo basale vel glabrum omnino; lobi calycis anguste trian-
gulares, 3—4 mm longi, marginibus glandularibus, abaxiale cum oe ees petala + circularia,

alba; stamina 10, ancheris ibis vel raro pallide roseis; carpelli et styli 2—3(—4). Fructus 8
mm diam., plerumque urceolatus, glaber, coccineus in juventute ee in maturitatem coloris

vini vel ee ea ie raro ee ecu si in conditione bona cum lobis calycis nN in-
. st

sig vel reflexis); pyrenae 2—3, dorsaliter sulcatae,
lateribus + + plais, paulo aie vel paulo erosis.

Bush, 3—G6 m tall; | year old twigs mid to dark brown; young shoots thinly
gee thorns stout, 2—4 cm long, shiny deep brown when young, re-
curved. Foliage deciduous, petiolate; petioles 1-2 cm long, thinly pubes-
cent, eglandular or with small black glands when young (as at Creston, B.C.);
blades 3.5—6 cm long, ovate to ovate-rombic, or less commonly broad el-
liptical to narrow-ovate, with 3—4 sharp lobes per side, margins finely double
toothed, 4—5 veins per side, somewhat impressed above, densely appressed
hairy when young above, this abrading with age, pubescent principally on
the veins beneath, somewhat coriaceous, glossy above especially when young,
expanding leaves reddish, the fall colour bronze. Inflorescence a domed panicle,
10—20 flowered, the branches variably villous, gland-dotted, with cadu-
cous linear, gland-margined bracteoles. Flowers 12-15 mm diam, notably
cup-shaped when young; hypanchium externally villous only at base or completely
glabrous; calyx lobes narrow triangular, 3-4 mm long, the margins glan-
dular, abaxially with a few hairs; petals + circular, white; stamens 10, an-
thers ivory or occasionally very pale pink; carpels and styles 2—3(—4). Fruit
8 mm diam., generally flask-shaped, glabrous, red when young but later
burgundy to deep purple (occasionally almost black), when in good condi-
tion with strikingly long erecto-patent calyx lobes but these sometimes shorter
or reflexed; nutlets 2—3, dorsally sulcate, their sides flat, a little plump, or
shallowly eroded.

Distribution.—W idely distributed and common from the Okanagan val-
ley of southern British Columbia and northern Washington to northern Idaho
(Fig. 6). This is an extraordinarily handsome hawthorn in late August and
early September.

Crataegus okanaganensis in similar to C. williamsii of northwest Montana as
illustrated by the numerical analysis (Fig. 1) but with more glossy foliage

Puipps AND O’KeNNoN, New species of Crataegus from Western North America 181

|-123° -120°| E \-415°

51°

oes. WA

N

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i ip
i}
I

N
bo
roe :
|-123° -12 ies le eos

Fic. 6. Distribution of C. okanaganensis J.B. Phipps and O’Kennon.

of a much more coriaceous texture and the color usually yellowish-green in
mid-season, the blades a little broader and the lobes usually less sharp, thorns
generally stouter, different anther color, and mature fruits deep purple (Fig.
9d) instead of sometimes blood or deep red; the young fruit is a brilliant
red (Fig. 9e) at the stage when C. wi/liamsii may be pale to deeper dull red.
The color change from brilliant red (late August) to a deep dull purple (late
September) is nearly unique among Crataegus species. In northern Idaho at
2300 ft, C. okanaganensis flowered in 1994 a good week before C. williamsii at
the same altitude in northwest Montana. Otherwise identical material with
five stamens is occasionally found. Crataegus okanaganensis was repeatedly
collected in the 1980s in Idaho by Fred Johnson though without recognition.

The two taxa may of ser. Purpureofructi be separated as follows:
1. Fresh anthers white or cream; inflorescence branches thinly to moderately

hairy; hypanthium usually glabrous; leaves coriaceous (Idaho and west).
okanaganensis
1. Fresh anthers bright pink; inflorescence branches densely hairy; leuk part

of hypanthium densely hairy; leaves thin (nw Montana). ........:.0eeee . williamsii
Additional specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Val-
ley, Westside Rd. NW of Vernon, ca. 2 km S of intersection with 97, 1450 ft, hedges on
both sides of road, 03 May 1994, Phipps. _ 6900 (UWO); ca. 1.2 mi E of ee on
road to Mabel Lake between 200 m & 25 m W of “Autobody collision & repairs” sign
1275 ft, hedgerow, bush, 3 m tall, fruit burgundy, 20 Aug 1994, Phipps, J.B. & O' Xenon,
RJ. 7016 (UWO); Rough grass with hawthorns on Hwy. 97 near entrance to Kelowna

182 Stipa 18(1)

airport, 5 side of Airport Dr., alt. 1800 ft, bush 6 m tall, wine red, — deep and mee
+ ellipsoid, 17 Aug 1994, Phipps, J.B. & O’Kennon, R.J. 6975 (UBC, UWO); ca. 8 km
of Vernon center at Burke’s Pick-your-own on Pleasant eve Rd., bes of ro ee
407 m; 5 m bush, dull burgundy, ellipsoid fruic, 24 Sep 1993, Phipps, J.B. & O’Rennon,
RJ. 6777 (BRIT, MONTU, UWO, WS); Shuswap Valley, just E of Enderby, across bridge,
S side of rd. on mound by roadside depression, alt. ca. 1250 ft, bush, 5 m tall, 10 stamens,
in bud, 05 May 1994, Phipps, J.B. 6912 (UWO), Okanagan Valley, ca. 9 km N of center of
Vernon off E side of 97, just S of jct. with 97A & just N of Baker Hogg Road, alt. 350 m,
hawthorn thickets to N side of long cleared ride, bush, 5 m tall, 10 white anthers, 03
eas B. 6894 (UWO); aries Valley, ca. 9 km N of center - Vernon off -
of 97, just S of jet. with 97A & just N of Baker Hogg Road, alt. 350 m, hawthorn
(mainly) eee to N side of long ee ride, bush, 5 m tall,10 white wees O03 May
1994, Phipps, J.B. 6894a (UWO); Shuswap Valley, ca. 2.8 km E of Enderby on Kingfisher
Rd., * hedgerow S side of road opposite house with wooden fence, N side, alt.1280 ft,
bush, 4 m tall, possibly = JBP & RO’K 6812, foliage somewhat reddish, stamens 10, cream
anthers, 0S May 1994, Phipps, A B. 6921 (BRIT. UWO); ca. 1.2 mi E of enn on road
to Mabel Lake between 200 m & 25 m W of “Autobody collision & repairs” sign
ft, ee bush, 5 m tall, ae reddish burgundy, 20 Aug 1994, Phipps eB; E0 ‘Kennon,
RJ. 7017 (UBC, UWO); Salmon Arm, large hawthorn fs asture just N of railroad and W of
ee ee section, alt. ca. 1900 ft, bush, 3.5 m tall, opposite building with blue
strip, flowers very cup-shaped, anthers 10, very pale pink. 11 May 1994, Phipps, J.B. 6966
(BRIT, CAN, UWO),; E side of 97A 0.5 mi N of intersection with 97, scrubby slopes with
long cleared area, bush, 5 m tall, scarlet fruit, 19 Aug 1994, Phipps, J.B. & O’Kennon, RJ.
7000 (UWO), Okanagan Valley, about 8 km NNE of center of Vernon, Pleasant Valley
Rd., behind Vowle’s residence, hedge at § boundary of Burke's PYO, alt. ca. 380 m, dense,
bread mature hedge, bush, 3 m tall, nearest top of hedgerow, 10 mnie ivory, 02 May
1994, Phipps, J.B. 6882 (DAO, UBC, UWO); Castlegar, near Selkirk College, left-hand
slope towards bottom of Rosedale Rd., alt. 970 ft, scrubby area, bush 2.5 m tall, prob.
JBP & RO’K 6826, stamens 10, anthers white, 06 peels 994, Phipps, J.B. 6929 (BRIT,
TRT, UBC, UWO); Okanagan Valley, Westside Rd. NW of Vernon, ca. 2 km § of inter-
section with 97, 1450 ft, hedges on both sides of road, iH 3 May 1994, Phipps, J.B. 6901
(UWO), Okanagan Valley, west side of hwy. at entrance to Kelowna airport, alt. 1800 ft,
open scrub, bush, 4 m tall, stamens 10, anthers ivory to palest pink, 05 1994, —
J.B. 6907 (BRIT, TRT, UBC, UWO); West side of hwy. 21, just N of, and in view of,
Canada Customs post, ae edge, 2200 ft, tall, 5 m bush, 27 Sep 1993 J.B. Phipps,
J.B. & O’Kennon, RJ. 0832 ID, UWO); NNW of Vernon on Westside Rd., ca. 2.5 km SW
of intersection of 97 and so hedges along roadside, back of field, W side of rd., bush, 5
m tall with purple shrivelled fruit, 24 Sep 1993, Gee & O’Kennon, ce 6788 (CAN,

—

TRT, UBC, UWO); E side of hwy. 21, ca. 250 mS of Creston Ferry Rd. to Hood Rd., this
species very common here, 1970 ft, dense bush, 3.5 m tall, see ped fruit, 27
Sep 1993, Phipps, J.B. & O’Kennon, RJ. 6831 UD, MO, UWO); E side of hwy. 21, imme-
diately S of jet. with BC 3, north of Creston, ale. 1850 ft, pee bank, 2.5 m tall bush,

fruic shrivelled, black, 27 See 1993, Phipps, J.B. & O’Kennon, RJ. 6830 (UWO); Castlegar,
sce of Campus Rd. off Frank Beinder Rd., grassy slopes with scattered hawthorns, bush,

, lvs sharply lobed, abundant burgundy fruit, 27 Sep 1993; Phipps, J.B. & O’Kennon,
: 7 6826 (BRIT, UBC, UWO, US); Larry Calder property, E of 97 about 9 km NNE of
Vernon, scrubby partly cleared hillside, 475 m, bush, 4 m, reddish foliage, fruit a good
purple, 26 Sep 1993, Phipps, J.B. & O’Kennon, RJ. 6819 (l IWO); Kalamalka Prov. Park, S
of Vernon, within 200 m of yellow gate, at parking lot, dampish depression, ca. 1375 ft,

Puipps AND O’KENNON, New species of Crataegus from Western North America 183

common, multistemmed bush, fruit oblong-orbicular, burgundy, 24 Sep oie ee SB.
= O’Kennon, R.J. 6798 (BRIT, UWO); NNW of Vernon on Westside Rd., ca. 2.5—3 km

of intersection ‘of 97 and 97A, hedges along roadside, back of field, 5 m ae with
ny fruit, 24 Sep 1993, Phipps, J.B. & O’Kennon, R.J. 6791 (UWO); E side of hwy. 21,
immediately S of jct. with BC3, north of Creston, scrubby banks, 1850 ft, 2.5 m bush,
leaves green turning purple, oe burgundy fruit, 27 Sep 1993, Phipps, J.B. & O'Rennon,
R.J. 6828 (UWO, V); N of Creston, E side of Hwy. 21, grassy banks with hawthorns on
top, just S of junction with Hwy. 3, alc. 1850 fr, bucks 2.5 m tall, stamens 10, anthers
white, equals ]BP & RO’K 6828, 07 May 1994, Phipps, J.B. 6930 (UWO), Salmon Arm,
large hawthorn pasture just N of railroad and west of central business section, alt. ca. 1900
ft, bush, 3.5 m tall, fruit bright red, equals JBP 6966, 20 Aug 1994, Phipps, J.B. G O’Kennon,
RJ. 7024 (BRIT, TRT, UWO), E side of 97A 0.5 mi N of intersection with 97, scrubby
slopes with long cleared area, a pa on south side of site, scarlet fruit, 19 Aug 1994,

Phipps, J.B. & O’Kennon, RJ. 700 WO), E side of 97A 0.5 mi N of intersection with
97, scrubby slopes with long ae area ate ee ant on south end of site, scarlet fruit,
19 Aug 1994, Phipps, J.B. & O’Kennon, RYT 2 (UWO); Hwy. 6 ca. 6 road mi W of

Fauquier, west of AoW Lake in Inookalin ane ie fenceline in farmland, alfalfa fields,
alt. 475 m, bush, 4 m tall, fruit bright red, 22 Aug 1994, Phipps, J.B. G O’Kennon, RJ.

7045a (UBC, UWO); Okanagan Valley NNE of Vernon, Pleasant Valley Rd., hedge be-

hind Vowle’s residence, E end boundary fence with Burke’s P-Y-O, alt. 380 m, bush 3 m

tall, fruit large, plump, purple-red, hairy, with stout erect calyx lobes, 19 Aug 1994, Phipps.
B. & O’Kennon, R.J. 6996 (UBC, UWO); Slocan Valley, S of Slocan, above swamp cross-

ing river along small road off Hwy. 6 & just N of Perry Siding, hedges and brushy slopes,
alt. 1775 ft, bush, 5 m tall, fruit bright red, 22 Aug 1994, Phipps, J.B. & O’Kennon, RJ.
7048 (UWO); Shuswap Valley, just E of Enderby, across bridge, E side ar road, 200 m N of
gate, alt. ca. 1250 ft, level open field, growing along roadside across from cattle feeder, 4
m tall, fresh ancth of , 15 May 1995, Donovan, L.S. & D. 2741 (UWO); Shuswap Valley,
just E of Enderby, across bridge, E side of road, alt. ca. 1250 ft, level open field, growing
along roadside across from cattle feeder, 2.5 m tall, fresh anthers cream, 15 May 1995,
Donovan, L.S. & D. 2744 (UWO); Okanagan Valley, Kelowna, service road at W side of
airport, alt. ca. 1250 ft, fenceline, bush 3 m tall; fruit brilliant red, ellipsoid, 17 Aug 1995,
Phipps, J.B. 7155 (UWO); Okanagan Valley, Oyama, ca. 0.5 mi E of E end of spit, alt. ca.
1320 ft, hedgeline on clay soil, bush, 4 m call, fruit re red, 17 Aug 1995, Phipps,
nmep el (UWO); Okanagan Valley, Oyama, ca. 0.5 mi E of E end of spit, alt. ca. 1320 ft,
hedg ‘lay soil, bush, 4 m tall, ‘fruit ellipsoid, 17 he 995, Phipps, J.B. 7159 UWO).
U. S.A. IDAHO. Bonner Co.: Lower Priest River, river bank, alt. 660 m., scarlet fruit,
07 Aug 1897, Leiberg, John B. 2857 (OSC); Pend Oreille R. across from Oldtown, N of
ee Section 24,T56N, R6OW, scrub just above high water mark, dominated by . evant
0 ft, 07 Aug 1986, Johnson, Fred 86126 (IDF); Leclede, Riley Creek, 8 mi R.
at a Oreille R. (Albenai Falls Reservoir) TSGN, R3W sec.30, riparian a nie res-
ervoir climax forest, gentle slope, common, 12 Aug 1988, Jobuson, F. 88079 (IDF); Pend
Oreille R. across from Oldtown, N of bridge, TSON ROW $24, 2000 ft, shrub fringe just
above high water mark, abundant, 07 Aug meas Fred 86126 (ID (#93728); 6 mi
E of Priest R, along Albenai Falls Reservoir in T , R4W sec. 34&35, riparian fringe,
1900 ft, common, 12 Aug 1988, Te F. 88082 “(IDE): Section 28,T56N, R3W along
Pend Oreille R. reservoir, pete ide fringe of trees/shrubs by hayfield, many thornless trees,
height to 15 ft, 07 Aug 1986, Johnson, F 86124 (ID, IDF). Boundary Co.: US 95, about
km S$ of jct. with Idaho | & S$ of turnout on right below tall conifers, alt. 2250 ft, on E
plowed field backed by steep rocky, treed slope, dense bush, 3 m tall, stamens 10, anthers

184 Stipa 18(1)

ivory, 08 May 1994, Phipps, J.B. 6932 ID, UWO); US 95, ca. 5 km S of jet. with Idaho |
& cee S of turnout on right below call conifers, alt. 2250 ft, on E ee field backed
steep rocky, treed slope, dense bush, 3 m tall, stamens 10, 07 May 1994, Phipps, J.B.
bone 3 (ID, UWO, WS); US 95 2 miS of jct. with Idaho 1, W side of road, ae 27 Sep
1993, Phipps, J.B. & O’Kennon, R.J. 6834 (UWO),; 2.6 mi § of jct. US 95 & Hwy. 1, in
TOAN, RIE sec.28 (SE of C ee roadside shrub fringe, 2200 ft, fae slope, 18 ft tall

x 22 ft wide, 5 main stems 3—5, possible escape, no farms near, 12 Aug 1998, Johnson, ie)
88072 (IDF). Kootenai Co.: floodpl ain of Spokane R. at Coeur d’ ieee 1/4 mi W of US
95, grassy meadow with scattered trees, no slope, open, 2000 ft, rare with C. dowglasii, 10

Jun am Johnson, FD. IDF (#8604 18)); Spokane River, 1/4 mi E of Washington border,
2100 ft, among granitic rocks and sand, ae to3m, Aug 1986, OED F. 8644 (ID
(#93 oy: Coeur d'Alene, 20 Jun 1930, t, JH. ), 20 Jun 1930; floodplain of Spo-
R. at Coeur d’Alene, 1/4 mi W of [ os grassy meadow with saniieted tree/shrubs,
elev. 2000 ft, 10 Jun re , Johnson, Fred ID (4937 30) re floodplain of Spokane R., at Coeur
d'Alene, 1/4 mi W of US 95, grassy sae with scattered trees & shrubs, no loge: open,
2000 ft, 16 Jun 1981, re ED. (DF). WASHINGTON. Okanogan Co.: Westside
Rd. ca. 7 km N of Tonasket, 975 ft, eee riverside, 7 m bush, smaller leaves, burgundy
fruit, 30 Sep —— Phipps, J.B. & O’Kennon, RJ. 6868 (UWO); Westside Rd. about 4 km
N of Tonasket, alt. 975 ft, bush 5 m tall, riverside hawthorn thicket with a few poplars,
fruit + erect, ee red, broad-elliptic, 23 Aug 1995, Phipps, J.B. 7175 (UWO, WS). Spo-
kane Co.: ie damp places, Swksdorf. WIN. 919 (WS), Aug 1889. Whitman Co.:
Pullman, Piper, C.V. (CAS), May 1902; Westend; 1800 ft, spring fae May, R. 91 (RMYV).

ENGLAND: ele Surrey, Kew, Royal Botanic Gardens; Kew #197-67, 19728; 22 May

1985, Phipps, J.B. 5651.

~ a
is
5

The discovery of C. okanaganensis and its clear relationship to C. wi/liamsi1
merit the creation of a new series, as follows:
Series Purpureofructi J.B. Phipps & O’Kennon, ser. nov. Type: C. okanaganensis
J.B. Phipps and O’Kennon.
Frutices, plerumque grandes, vel arbores parvae; ies ie mediocres i in longitudine (2—

cm longae). Folia plerumque + coriacea (praeter C. wiflian laminae plerumque oe.
t |

7

3—4 brevibus patentibus per
lacus. Inflorescentia 10-25 nae: Flores denon 15 min) crane stamina 5-10; carpelli
A

(vel eliptico-oblongae vel rhombo-ovatae), Se ace lobis

et styli 2—4. Fructus cineus, plerumqu 1eus in maturitate
Septembri exeunte fierens oe ae vel atropurpureus, orbic mye vel ampulliform mis; pyrenis
34, lateribus erosis vel planis.

Bushes, usually large, to small trees; thorns medium in length (2—4 cm).
Leaves generally + coriaceous (except C. wi/liamsi?), blades usually + rhom-
bic (to elliptic-oblong or rhomb-ovate), usually with 3—4 short outward-
pointing lobes per side. Inflorescences 10-25 flowered; flowers not large
(10-15 mm diam.); stamens 5—10; carpels and styles 2—4. Fruit red, usu-
ally brilliant red in late August, becoming deep paaekia to dark purple
at maturity, orbicular to flask shaped; pyrenes 3—4, sides eroded or plane.

Distribution.—A distinctive series with two known species and perhaps
a couple of others found from the interior Pacific Northwest and southern
British Columbia to northwestern Montana. Crataegus wiliamsii was origi-

PHipps AND O’KENNON, New species of Crataegus from Western North America 185

nally placed in series Rotundifoliae by Eggleston but the ellipsoid fruit, long

calyx lobes in fruit, dark-colored twigs and somewhat erose nutlets suggest

a different affinity. Our numerical analysis (Fig. 1) indicates no relation-

ship whatever to C. chrysocarpa in ser. Rotundifoliae.

3. Crataegus phippsit O’Kennon, sp. nov. (Fig. 7). Type: CANADA. BritisH
Co.uMBIA: Okanagan Valley, Pleasant Valley Rd., ca. 8 km NNE of Vernon center,
bush, 4 m tall, fruit red, 24 Sep 1993, Phipps, oe a and O’Kennon, RJ. 6780 (HOLO-
ryPe: UWO); tsoryres: BRIT, CAN, TRT, US,

Arbor vel frutex eae poe malta, lomi 1 alig reuli
panun purp /-bru inne , VELETES atrocinenerel; inj tut

>
1 breviter

li p ainébnuinneas ve | nigrae, leviter recurvatae, aliquantu
tenues, 2-4 ems longae. Folia decidua, petiolata; petioli 1.5—2.0 cm longi, fies
laminae 4-8 cm longae (sed ad 10 cm vel plus in blastis elongationis), late ellipticae vel
ovatae vel deltoideo-ovatae, lobis fere 3 + vadositer obtusis vel late angulatis per latus,
marginibus breve dentatis; ca. 4 (-5) venis secondariis per latus, ee breviter appresso-
pubescentes subter breviter pubescentes praecipue in venis, aliquantum coriaceae postea.
Inflorescentia panicula aaa 6-12 florata; ramuli dense es bracteolati;
bracteoli lineares glandulo-marginati. Flores 15-22 mm diam.; hypanthium externe breviter
tomentosum; obi pie late triangulares, ca. 5 mm longi, abaxiale pubescentes praecipue

inibus pe laciniatis; petala + circularia; stamina 10, antheris roseis
pallide: ee aie stigmatis capitatis. Fructus 12 mm diam., breve-pubescentes,
maturentes ab coccineo ae oan eo-purpureum vel nigrum, + apiece vel late urceolati,

obis calycis reflexis; pyrenae 3, vadositer sulcatae dorsale, lateribus planis.

‘Tree or large bush, S—6(—7) m tall, with main trunk, but sometimes suckering;
1 year old twigs dark purple-brown, older dark gray, young shoots densely
short pubescent; thorns somewhat sparse, deep brown to black, slightly recurved,
somewhat thin, 2-4 cm long. Foliage deciduous, petiolate; petioles 1.5—
2.0 cm long, pubescent; blades 4-8 cm long (but on shoots of elongation
to 10 cm or more), broadly elliptic to ovate or broad deltoid-ovate, with
about 3 + shallow rounded to broadly angled lobes per side, the margins
short-dentate; about 4-5 secondary veins per side, shortly appressed hairy
above, shortly hairy especially along the veins below, somewhat coriaceous
later. Inflorescence a 6—12-flowered flattened panicle, the branches densely
short-tomentose, bracteolate, the bracteoles linear, gland-margined. Flow-
ers 15—22 mm diam; hypanthium externally densely tomentose; calyx lobes
broad-triangular, ca. 5 mm long, pubescent abaxially, especially below, the
margins glandular—laciniate; petals + circular; stamens 10, anthers pale pink;
inside of hypanthial cup mostly glabrous, bristly at center; carpels and styles
2-4, the stigmas capitate. Fruit 12 mm diam., short-hairy, ripening through
shades of red to plum-purple to black in colour, + orbicular to broadly flask-
shaped, pubescent, with reflexed calyx lobes; nutlets 3, shallowly dorsally
ridged, lateral walls + plane.

Distribution.—This species is not so common as the other two described

186 Sipa 18(1)

\ Lee

Yaa ys
MS) i772
AWAS

2 34 TF
nae

Fic. 7. Line drawing of C. phippsii O’Kennon. Inflorescence flowers and parts from Phipps
6891 (UWO); infructescence, fruit and parts from Phipps 6780 and O’Kennon (UWO), two
smaller isolated leaves from Phipps 6780 and O’Kennon (UW O), magnified leaf (bottom left)
from Phipps 6891 (UWO). Scale bars = 1 cm.

PHipps AND O’KENNON, New species of Crataegus from Western North America 187

in this paper but is nevertheless widespread and has been found at fourteen
localities ranging from the Okanagan Valley of southern British Columbia
and northern Washington to the Flathead Valley of Montana (Fig. 8). We
have never found it numerous at any site.

Crataegus phippsii is one of the most distinctive species of hawthorn to
have been found in North America in recent decades and does not appear
to be closely similar to any other known in the west, a feature emphasized
in Figure 1. It is a large hawthorn, reminiscent of C. mo//zs, the latter a common
species of the interior of the continent, in foliage, habit and pubescence while
in its red (Fig. 9a) changing to purple (Fig. 9b) to black fruit it is similar to
a number of mostly previously undescribed species of the Pacific North-
west and British Columbia. The foliage colours attractively in autumn (Fig.
9b, Of). The mature bark sometimes flakes off in longitudinal strips, an unusual
feature in hawthorns (Fig. 9c).

Crataegus phippsii was first collected by B.T. Butler (nos. 527—523) in
1908 and these specimens were cited by Eggleston (1909) in his protologue
for C. williamsit. The many differences from C. wi/liamsii, however, show
that C. phippsii in not in this affinity. That Eggleston thought it somewhat
distinct is shown by his annotations on the sheets of “var. ovata,” a name,
however, never published. Beyond this brief glimpse of notoriety, the present
species, very striking to the present authors, remained totally overlooked
until our collections of recent years.

The large, broad leaves with rounded lobes and hairiness of all parts sug-
gests an affinity with series Mo//es. Superficially, it most nearly resembles
the local species C. moelensis Sarg. of Missouri, a taxon which, however, has
not been seen for many years. A combination of stamen number, anther color,
foliage size and shape, and fruit color are the main characters distinguish-
ing the species in the Mo//es group as shown in the following key. For con-
venience, I am including C. greggiana in this key although it is now placed
in ser. Greggianae.

KEY TO C. PHIPPSII AND MEMBERS OF SERIES MOLLES

1. Foliage small, on short shoots mainly 4 cm —
2. Flowers 1-1.5 cm diam.; stamens Les 10 C. greggiana
Flowers ca. 2 cm diam.; stamens 2 C. lanuginosa
I. Boras. je on short ie eal 5—8 cm long.
3. Anthers pink.
4. Stamens 5—10

5. Fully ripe fruit orange-red; leaves + unlobed. C. noelensis

5. Fully ripe fruit deep purple to black; leaves evidently lobed. ........ C. phippsii
4. Stamens 20.

6. Fruit red. C. texan

6. Fruit yellow. C. viburnifolia, C. kellogii

3. Anthers white to cream.

188 Sipa 18(1)

{ 1.423° '_490° "4 12° "444°
\
x

| l l
\
I !
1 MT
WA 1 ff
1 N
{ \
+
| \
fi: ee \
i a a
ae * ID 7
\ se
rf a
Re f'28° io er [fits
Fic. 8. Distribution map of C. phipps7i O’Kennon.
7. Stamens 20. C. mollis
7. Stamens 10. C. submollis, sens. lat.

Additional Specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Val-
ley, 8 km NNE of center of Vernon, Pleasant Valley Rd., behind Vowle’s residence, hedge
at S boundary of Burke's PYO, dense, broad mature hedgeline, bush 4 m, commencing
ee -8 pale pink anthers, 02 May 1994, Phipps, J.B 6878 (BRIT, CAN, TRT, UBC,

O, US); Okanagan Valley, 8 km NNE of center of Vernon, Pleasant vey Rd., be-
i ee s residence, hedge at S boundary of Burke’s PYO, dense, broad mature hedgeline,
bush 4 m, foliage glossy, pedicels hairy, 10 pink anthers, 02 May 1994, Phipps, J.B. aed
(CAN, TRT, UBC, UWO); Okanagan vay scrubby slope above dirt road N
bound into O’ Keefe Ranch, opposite L Cross Rd. off 97A, alt. 1290 ft a.s.l., stamens
10, anthers light pink, equals JBP & RO’K 6803, 03 May 1994, Phipps, J.B. 6891 (UBC,
UWO, WS); ca. 10 km N of Vernon center, on dirt road N-bound into O’Keefe — off
Hwy. 97A at L.& A. oe Rd., dry rocky hillside, suckering clump of trees to 5 m
across, bark coarse, plated, peeling, fruit purple, 25 Sep 1993, Phipps, J.B. & oe
RJ. 680 3 (CAN ae is Pleasant Valley Rd. ca. 8 km NNE of Vernon Center, southern
boundary hedge oe Burke’s PYO behind Vowles’ residence, alt. ca. 380 m, dense, broad
mature hedgeline of Crataegus, bush 4 m tall, fruit deep red, 24 Sep 1993, Phipps, J.B. G
O’Kennon, R.J. 6780 (BRIT, CAN, MO, UWO); ca. 5 km NNE of Enderby, on E-W rd.
between Shuswap R. and Hwy. 97A, long hedgerows, alt. ca. 1375 ft, bush, 7 m tall, fruit
red, 21 Aug 1994, Phipps, J.B. & O’Kennon, RJ. 7038 (UWO), ca. 3 km NE of Enderby on
road to Sicamous (Hwy. 97A), bottom of scrubby slope on east side, alt. ca. ve ff as
of 5 m bushes, reddish fruit, 21 Aug 1994, Phipps, J.B. & O’Kennon, RJ. 7037 (DAO, U
ca. 3 km ENE c of Armstrong on E-W aligned rd. just ENE of A Armstrong airpot cs ate
S side of road, alt. 1450 ft, bush 5 m call, 21 Aug 1994, Phipps, J.B. & O’Kennon, R.J. 7041

fal

PHIpPs AND O’KENNON, New species of Crataegus from Western North America 189

Fic. 9. Various pictures of C. phippsii O’ es and Cc. onan B. Phipps & O’Kennon:
Aippsii, northern Okanagan, BC, 19 Aug

1995; b) typical later September fruit ae of C. phippsti _. fall foliage color beginning,

a) typical reddish late suguse fruit color

northern Okanagan, BC, 24 Sep 1993; c) trunks of C. pAzppsii showing shredding bark,
shores of Palmer Lake, eae late Aug. 1994; d) full ripe fruit color of C. okanaganensis,
Westside Rd., nr. Vernon, BC, 24 Sep 1993; e) late August fruit color of C. okanaganensis,
nr. Kelowna, BC, 17 Aug 1995; . R.J. O’Kennon holding fruiting branch, Palmer Lake,
Okanagan Co., WA, 01 Oct 1993; note fall foliage

190 Sipa 18(1)

(UWO); ca. 6 km N of Armstrong, branch of Deep Creek, thickets, primarily C. —
n narrow flood- om alt. 1450 ft, tree 4 m call, fruit red, 21 Aug 1994, Phipps, J.B.
O’Kennon, R.J. 7034a (UWO), ca. 3 km NW of Armstrong, N-S road parallel to ee

Rd. and to the 7c - it, E side, alone by telephone pole, alt. ca. 1400 ft a.s.1., bush 4
tall, dark green leaves, dull reddish fruit, 21 Aug 1994, Dbinps J.B. & O'Kennon, RJ. 7033
anes er UWO).

MONTANA. Flathead Co.: about 0.5 km along Tower Rd. off US 93, near SW
corner Blachead Lake, hawthorn thicket along trail, rear of small field, alt. 2950 ft, equals
JBP 7078, bush 7 m tall, eee saa all others, at popcorn, pace pink, 20 May 1997
Phipps, J.B. & O’Kennon, RJ. 7591 (MONTU, UWO). Lake Co.: Hwy. 93, mipost 63.4,
Tower Rd., (major pablillo), 2. if mi. NW of Polson Bridge, 8— a 12)-14 flowers, 10 pink
anthers, leaves hairy, shaggy, in green bud stage (forced), 18 May 1995, O’Kennon, RJ. 13156
(UWO); Flathead Lake, SW border, 29 Aug ee Butler, B.T. 522 (NY); Rocky cs
Road NW of Polson at 1.7 mi N of hwy 93 and 0.3 mi NE of Jim’s Road, alt. 292(
extensive thickets near lakeshore behind new ore bush, 5 m tall, fruit orbicular, —
dening, 25 Aug 1994, Phipps, J.B. & O’Kennon, R.J. 7080 (BRIT, ID, MO, MONTU, UWO),
ger Lake, SW border, 19 Aug 1908, Butler, B.T. 523 (NY); Flathead Lane, off Tower

Rd., off hwy 93 few mi NW of Polson, thickets along creek back of field, alt. 3050 ft, tree
Sm ih “fea partly deep partly paler, 25 Aug 1994; Phipps, J.B. & O’Kennon,
R.J. 7078 (DAO, MONTU, UWO, US); Flathead Lake, SW border, 19 Aug 1908, Butler,

Ba, O21 NY) ano Olianogan Co.: Palmer Lake, N side., open hawthorn
thicket at upper level of pebble beach at picnic spot/campsite, just E of Chopata Lodge,
small tree, 6 m tall, mostly finished flowering, stamens 10, anthers pale pink in bud,=JBP
& RO'K 6874, 04 May 1994, Phipps, J.B. 6904 (ID, MO, UWO, WS); W of US 97 on
crossroad at Ellisford, in floodplain to S side of road, bush, 7 m tall, anthers 10, pale pink,
equals JBP & RO'K 6876, 04 May 1994, Phipps, J.B. 6906 (US, UWO, WS); Palmer Lake,
N side, open hawthorn thicket at upper beach level on campsite just E of Chopata Lodge,
alc. 1150 ft, fine tree, 7 m tall, burgundy fruit; 30 Sep 1993, Phipps, J.B. & O’Kennon, RJ.
6874 (DAO, ID, MO, We WS; E shores of Palmer Lake, 0.7 mi S of campsite at N end,
thickets, alt. 355 m, bush, 4.5 m tall, deep plum-red fruit, 28 Aug 1994, Phipps, J.B. &

O sui R.J. 7108 (BRIT, ID, UWO, WS); Okanogan Valley, cross-road at Ellisford, alc.
950 ft, bush 7 m tall in field in floodplain, S side of road, fruit blackish, 01 Oct 1993,
si a & O’Kennon, RJ. 6876 (UWO, WS).

ACKNOWLEDGMENTS

We wish to thank the National Research Council of Canada for supporting
this research, Susan Laurie-Bourque of Hull, Quebec for the line illustra-
tions and Antony Littlewood of the Department of Classical Studies, The
University of Western Ontario, for checking the Latin diagnoses. Peter Wood
of Selkirk College, Castlegar is thanked for field assistance. J.B. Phipps thanks
R.J. O’Kennon for supporting the color printing.
REFERENCES
EGGLESTON, W.W. 1909. New North American Crataeg/. Bull. Torrey. Bot. Club 36:639—
642.
Hoimcren, N.H. 1997. Rosaceae. - A. Cronquist, et al., eds. Intermountain Flora, vol.
3A. New York Botanical Garden

n3
Pups, J.B. 199x. Taxonomic and eh atural problems in black and burgundy-fruited
Crataegus (Rosaceae) species: C. erythropoda, C. rivularis and C. saligna. Sida, submitted.

Puipps AND O’KENNON, New species of Crataegus from Western North America 191

Puipps, J.B. 1998. Introduction to the red-fruited hawthorns (Crataegus, Rosaceae) of western
North America. Canad. J. Bot. 76(in press).

SARGENT, C.S. 1907. The black-fruited hawthorns of western North America. Bot. Gaz.
44:64—66.

WesH, S.L. 1987. Rosaceae. In: A Utah Flora. Great Basin Naturalist Mem. 9:520-543.

192 Sipa 18(1)
BOOK REVIEW

ANNE HutTcHINGs, and ALAN HAXxTON Scott, GILLIAN Lewis, and ANTHONY
CUNNINGHAM. 1996, Zulu Medicinal Plants: An Inventory. ISBN
0-86980-893-1, pbk; ISBN 0-86980-923-7, hbk). University of Na-

tal Press, Private Bag XO1, Scottsville 3209, Republic of South Af-
rica, in association with University of Zululand and National Botani-
cal Institute. Exclusive distributor: International Specialized Book Services,
Inc., 5804 N.E. Hassalo See Portland, OR 97213-3644, U.S.A.
(503) 287-3093; FAX (503) 280-8832. $110.00. 450 pp.

The subcitle understates the amount of information in this tightly organized compendium

of information about plant used by Zulu traditional healers. The authors have chosen to

arrange their entries in phylogenetic order to facilitate comparisons ae related plants.

As this publication deals exclusively with plants of southern Africa, the classification fol-
lows that used in the National Herbarium. Genus numbers reflect the standards used in
herbaria arranged in the Englerian system with additional standards for ferns and grasses.
These are referenced in the introduction

The entry for each species includes geographical distribution, a short description, local
se by Zulu healers and

c

names in English, Afrikaans and Zulu followed by documente
then those referring to use by other groups in southern Africa. iysiolorica effects, Chemical

constituents and biological properties are added where information is available. These gaps
will tease and stimulate further research conducted, it is hoped, under the ethical and legal
codes protecting intellectual property rights

rere is a wealth of information not ae on the medicinal uses of these plants, but by
the social and cultural uses too. These range from love potions to charms against lightning
and incense to invoke the goodwill of ancestors. However, the authors do warn readers not
to experiment!

Por those who do not own a copy of Watt and Breyer-Brandwijk’s 1962 edition the Me-
dicinal and Poisonous plants of Sonthern and Eastern Africa this book is a must. For those for-
tunate to own a copy this will be a valuable supplement, for at least a third of the 1032
species mentioned here are new.

With the worldwide renewal of interest in natural products this book has a huge poten-
tial readership. In each foreword, we are reminded of the value, and ephemeral nature, of
traditional knowledge. This book makes it less likely we will lose the traditional ethnobo-
tanical knowledge of the Zulu nation.—Fiona Norris

Sipa 18(1): 192. 1998

TAXONOMIC CLARIFICATION OF ATRIPLEX
NUTTALLIT (CHENOPODIACEAE)
AND ITS NEAR RELATIVES

HOWARD C.STUTZ
Department of Botany and Range Science
Brigham Young University
Provo, UT 84602, U.S.A.
STEWART C. SANDERSON
USDA Forest Service Shrub Sciences Laboratory
Provo, UT 84601, U.S.A.
ABSTRACT

In 1874 Sereno Watson described Atriplex nuttallii as a new species, citing his number
981 collection, made in meee from northern Nevada, as a representative. His collection
t 1 from a separate population all mounted on one sheet

consis sted of th three p

is A. canescens, two are the new species, A. nutral/i. Failure to recognize the arena
collection and collection site of A. nitteallié hie resulted in assignment of the name A. nuttallit
to several different Atriplex species in western North America. However, by using quanti-

tative as well as Srilteaee differences, A. nuttallii is readily diy acura from its near
relatives A. cuneata, A. falcata, A. gardneri, and A. tridentata. The principal distinguishing
features include plant size and habit, leaf dimensions, fruiting-bract size and shape, fruit-
ing bract appendages, chromosome number, saponin content, flavonol content, dates of
anthesis, and geographic distribution.

RESUMEN

En 1874 Sereno Wat lescribid Arripl tallt pecie, citando su recoleccién
numero 981, hecha en el norte del estado de Nevada, E.E.U.U., como representativa. Consiste
esta recolecci6n en tres a de distintos en pero colocudas todas en un mismo a
de herbario. Una de ellas e canescens y dos de la nueva especie, A. nuttalli. El n
darse cuenta los botanicos de hs ae ta original y su nase de origen ha tenido como consecuencia
la aplicacién del nombre de Atriplex nuttallii a varias otras especies de Arriplex del oeste de

orte América. Sin embargo, usando tanto caracteristicas Cuantitativas como cualitativas,
se distingue facilmente A. nuttallii de las especies afines, A. cuneata, A. falcata, A. gardneri,
y A. tridentata. Las CANCTE TsSIGes ae fos uniligad son la altura de la planta, es dimensiones

de la hoj re la forma y tamafio de ap éndices, el
contenido de saponinas, contenido de flavonoides, fechs de eantesis, y la distribucién peoples:
INTRODUCTION

In 1874, Sereno Watson described Atriplex nuttallii as new (Watson 1874)
and listed his collection number 981, obtained in 1868 in northern Ne-
vada, as a representative of the species. This collection consists of three plants

Sipa 18(1): 193-212. 1998

194 Stipa 18(1)

collected at different times in separate locations in northern Nevada, all
mounted on one herbarium sheet (GH!) (Fig. 1), and each labeled by Watson
as Obione canescens Moq. The plant on the right is A. canescens (Pursh) Nutt.
collected in June, 1868, in Unionville Valley, Pershing County, Nevada.
The other two are A. nuttalliz. The middle plant, which appears to have
been broken and folded back, was collected in September, 1868, in Thou-
sand Springs Valley, Elko County, Nevada. The plant on the left was col-
lected in July 1868 in Reese Valley, Lander County, Nevada. Currently there
are still extensive populations of these species in the valleys where Watson
collected them.

As shown in Figure 1, several botanists have recognized differences be-
tween these three plants and have provided annotations for them. In 1951,
Grant D. Brown annotated the plant on the right as Atriplex canescens (Pursh)
Nutt. In 1962, C.A. Hanson annotated the plants on the left and right as
Atriplex bonnevillensis Hanson and the central one as Atriplex falcata (Jones)
Standley. In 1972, I.J. Bassett and C.W. Crompton annotated the plant on
the right as Atriplex canescens, the central one as A. nuttallit var. falcata, and
designated on the annotation label, the left specimen as the lectotype of
Atriplex nuttallii &. Watson. Later, McNeil et al. (1983) designated as the
lectotype of A. nuttallii, a specimen collected in Saskatchewan, Canada, by
Bourgeau (s.7.), in 1857, and included by Watson among 10 other collec-
tors of A. nuttalliz, in addition to Watson 981. However, as directed in the
Senne Code of Botanical Nomenclature, Article 9.9 (Greuter et al.

94), —” when the material designated as type is found to belong to more
ve one taxon, a lectotype — may be designated,” and, as described in
Article 7.5 of the 1988 edition of the code (Greuter et al. 1988), “A lecto-
type is a specimen or illustration selected from the original material to serve
as a nomenclatural type when no holotype was designated at the time of
publication —. When two or more specimens have been designated as types
by the author ... the lectotype must be chosen from among them.” Since
Watson listed in the protologue, his number 981 as a representative of the
new species, he ostensibly considered it typical and, since this collection
consists of three specimens, one of them must be chosen as the lectotype.
Consequently, the lectotype of Atriplex nuttallii is the specimen on the left
(GH) as annotated by Bassett and Crompton in 1972. The designation by
McNeil et al. of a specimen collected in Saskatchewan, Canada as the lecto-
type of A. nuttallii, was therefore unnecessary. The illustration in Bassett et
al. (1983) of the Saskatchewan plants, is representative of a diploid A. gardneri
(Moquin-Tandon) Hall & Clements which is common throughout south-
ern Alberta, southern Saskatchewan and northern Montana but very differ-
ent from A. nattallii of northern Nevada. In his protologue, Watson de-

STUTZ AND SANDERSON, Atriplex nuttallii and its near relatives

1D

LELTD Ty Pe

*

ug ,
Asmotarat by td. Benn -
Department of Agriculture, Orta

SDENTINIED AY THE HERBARIUM OF
POMONA COLCece
CLAMEMONT. CaLironmia

Rasms Wass, Catiector,

Fic. 1. Atriplex nuttallii. The lectotype seis nuttallit (Watson 981, upper left corner)

collected in Reese Valley, Lander County, Nevada,

868. The central element is A.

cata cole in Heusane puns Valley ue eee Nevada, September, 1868.

va

Soe June 1868.

n Unionville Valley, Pershing County,

196 Sipa 18(1)

scribes A. nuttallii as being 1—2 feet high. This is the stature of the A. nuttallii
plants in the populations he collected in northern Nevada (Table 1) but not
A. gardneri plants which are mostly 6-12 inches tall, nor A. canescens plants
which are seldom less than 30 inches in height. The Saskatchewan plant
illustrated in Bassett et al. (1983) appears to be ca 12 inches in height.

In the botany treatment of the King expedition (Watson 1871), Torrey
provided identification for Watson’s collections and Watson provided the
descriptions (footnote, page 287). Referring to Watson 981, Torrey identi-
fied it as Obione canescens Mog. and Watson applied that name to each of the
three plants. Watson’s description of the collection specifically referred to
his number 981 and indicated that “most of the specimens have nearly or
quite wingless fruit. Others have the bracts considerably dilated, though
still less than is frequently the case (982),” indicating that he recognized it
as being anomalous, quite unlike other Obione (Atriplex) canescens plants. As
shown in Figure 2, there is considerable variation in the fruiting-bract char-
acteristics of plants in the Reese Valley population, with none being truly
4-winged.

Since Watson, following Torrey’s identification, labelled each of the three
specimens (Watson 981), Obione canescens, he apparently assumed the varia-
tion he saw in the populations in northern Nevada to be representative of
the variation displayed by Atriplex shrubs throughout western United States.
This is confirmed in his later description of A. nuttallii (Watson 1874) wherein
he lists Atriplex canescens Nuttall, not of James, Obsone canescens Mogq., and
Atriplex Gordoni Hook. as synonyms and gives its distribution as “from the
Saskatchewan to Colorado and Northern Nevada.”

This broad interpretation by Watson appears to have been the initiation
of a series of misconceptions concerning the geographic distribution of A.
nuttallii that we interpret to be confined to northern Nevada and north-
western Utah (Fig. 3). Standley (1916) listed the type locality of A. nuttallii
as “on the denuded hills of the Missouri River, about 15 miles below the
confluence of the White River, South Dakota.” Since this is the location
given by Nuttall (1818) as the type locality of Atriplex canescens (Pursh) Nutt.,
Standley clearly confused the two. Furthermore, this locality for Atriplex
canescens (Calligonium canescens Pursh) invites correction because its origin
given by Pursh (1814) is “in the plains of the Missouri River, near the Big
Bend” w
River.

Standley (1916) also mistakenly listed the distribution of A. nuttallii as

“alkaline plains and hillsides, Manitoba and Saskatchewan to northern Utah,

—

aich is several kilometers upstream from the confluence of the White

Colorado and western Nebraska” without mentioning Reese Valley and Thousand
Springs Valley in northern Nevada.

STUTZ AND SANDERSON, Atriplex nuttallii and its near relatives 197

Fic. 2. Fruiting-bracts of six randomly selected plants of Atriplex nuttallii from a popula-
tion in Reese Valley, ca 1 km west of Battle Mountain, Lander Co., Nevada. Each row across

represents one individual. Bar = 15 mm.

198 Sipa 181)

a

es:

Fic. 3. Distribution of known populations of Atriplex nuttallii.

en)

Hall and Clements (1923) likewise failed to include the type locality of
A. nuttalli in their description of the distribution of A. nvttallii as “Saskatchewan
and South Dakota to western Nebraska, New Mexico, Arizona, northwest-
ern California, eastern Washington, and Alberta.”

Recently, Basset et al. (1983) cited the distribution of A. nuttallii as “from
the Peace River District of Alberta to Manitoba. In the United States specimens
have been found as far south as Colorado.” Clearly this does not refer to the
A. nuttalli described by Watson from Nevada.

The failure to identify the original collection site of A. nuttallii in all
references has caused considerable misunderstanding of its identity and has
resulted in the assignment of this binomial to most of the suffrutescent shrub
species of Arriplex in western North America. Ulbricht (1934) listed A. nuttallii
S. Wats., A. buxifolia Rydb., A. cuneata A. Nelson, A. eremicola Osterh., A.
falcata Standley, A. gordonii Hook., A. neomexicana Standley, A. oblanceolata
Rydb., A. pabularis A. Nelson and A. tridentata Kuntze as synonyms of Obione
gardneri Mog. (Atriplex gardneri {Moq.] Standley). Hall and Clements (1923)
treat A. buxifolia, A. cuneata, A. eremicola, A. falcata, A. gardneri, A. pabularis
and A. tridentata as subspecies of nuttallii.

Standley (1916) listed eight species as close relatives of A. nuttallii (Se-
ries Nuttallianae): A. buxifolia, A. cuneata, A. falcata, A. gardneri, A. neomexicana,
A. oblanceolata, A. pringlei Standley, and A. tridentata. As suggested by Hall

Stutz AND SANDERSON, Atriplex nuttallii and its near relatives 199

and Clements (1923), A. pringlei appears to be an insignificant variant of
A. acanthocarpa (Torr.) S. Wats and A. oblanceolata is non-distinguishable
from A. gardneri. A. neomexicana is a common reoccurring hybrid between
A. cuneata and A. confertifolia (Torr. & Frem.) 8. Wats. (Hanson 1962) and
A. buxifolia is synonymous with A. gardneri. The remaining four species, A,
cuneata, A. falcata, A. gardneri, and A. tridentata, were treated by Hall and
Clements (1923) as subspecies of A. nuttallii. Since they and A. nuttalli are
clearly distinct phenotypically (Pope 1976, Tables 1, 2, Fig. 4) and geo-
graphically (Figs. 3, 5-8) we consider them best treated as separate species
as proposed by Standley (1916).

Because Watson listed A. gordoni (A. gardneri) as a synonym of A. nuttallis,
Hanson (1962) argued that the epithet zmftallii was superfluous and hence
illegitimate and that A. gardneri was the correct name for Watson's A. nuttallit.
This view was later accepted by Pope (1976) Stutz (1978), Stutz et al. (1979),
and Welsh (1984). However, as noted by McNeil et al. (1983), the listing
of A. gordoni (A. gardneri) as a synonym of A. nuttallit was accompanied by
a question mark and is therefore not a legitimate synonym and, according
to the Code, Article 55.2, note |, (Greuter et al. 1994) the epithet nattalli
is therefore not nomenclaturally superfluous.

As discussed by McNeil et al. (1983) the other synonyms of A. nuttallii
listed by Watson: (Atriplex canescens Nuttall, and Obsone canescens Moq.), are
also not legitimate synonyms.

MATERIALS AND METHODS

To better define A. nvttallii and its near relatives, herbarium specimens
of the suffrutescent Arriplex species of western North America were exam-
ined in several herbaria (BRY, CAS, GH, MO, NY, PH, RM, RSA, US, UC)
and specimens were collected and studied from the original collection sites
of each of the species included in the study (except A. gardneri for which
the exact location is not known). In addition, morphological measurements,
saponin production, flavonoid content, and cytological studies were made
of plants in several populations of each species. The morphological mea-
surements included fruiting-bract characteristics, plant height and width,
and leaf length and width. Chromosome counts were obtained from plants
in several populations of each species (Figs. 5—8). In some cases the counts
were obtained from root tips squashed in aceto-carmine stain, but most were
from aceto-carmine squashes of pollen-mother-cells derived from anthers
fixed in 5% acetic acid and stored under refrigeration. Saponin content was
determined by hemolysis of red blood-cells as described by Sanderson et al.
(1987). Flavonoid content was determined by paper chromatography as described
by Sanderson and Stutz (1984).

Taste 1. Plane and leaf characteristics of Atriplex nuttallii and its near relatives. Measurements were obtained from 20 plants in each population. Diploid A. cwneata
is not included because of extensive between-population variation. N = number ee en Data = mean with coefficient of variation in parentheses. Values
in each column with the same letter are not significantly different (p<.05).

Plant Leaf
Taxon N Height (cm) Width (cm)* Volume (dm°*)> Length (mm) Width (mm) I/w
A. nuttallii(6x) 9 32.6(0.41)* 47.4(0.39)4 101.5(0.94) 24.1(0.33)4 4.0(0.21)® 6.0(0.22)*
A. cuneata(Ax) 3] 12.8(0.31)8 51.2(0.33)4 41.3(0.88)48 20.9(0.22)4 8.3(0.28)4 2.6(0. 19)BC
A, falcata(2x) 12 12.9(0.44)8 24.6(0.31)8 9.4(0.74)8 23.7(0.58)4 3.6(0.41)8 6.5(0.23)4
A. gardneri(2x) 6 9.8(0.5.2)8 48.9(0.11)48 25.2(0.76)48 19.9(0.09)4 3.8(0.21)8 5.4(0.11)4P
A. gardneri(4x) 8 10.9(0.59)8 49.8(0.44)4 42.3(1.45)48 19.4(0. 19)4 5.3(0.28)8 3.8(0.18)B
A. tridentata(6x) 13 23.9(0.38)* 41.8(0.46)48 63.8(1.25)48 28.7(0.53)4 4.1(0.33)% 6.79(0.28)4

“Width of A. tridentata does not include root sprouts.
> Volume = height < (width)?

O0C

(L)81 Vals

TABLE 2. Fruiting-bract characteristics of Atriplex nuttallii and its near relatives. Measurements were made on 20 fruits from several randoml y selected avout in

each sampled population. Diploid A. cwmeata is not in because of extensive between-population variation. N = number of populations. Data = mean with
secant of variation in parentheses. Values in each er with the same letter are not significantly different (p<.05).
Fruiting-bract
Taxon N Length (mm) Width (mm) Uw Volume (mm/?)* Apex? No. of Appendages‘*
Terminal Teeth
A. nuttallii(6x) 10 4.5(0.10)® B0(O0A)ee) ~ iso 4A yee $9:0(0.15)" 2.4(0.07)8 3,2(0.24)8 2.0(0.14)®
A. cuneata(Ax) 15 6.1(0.16)" 5.0(0.22)4 1.3(0.14)© 165.3(0.59)4 1.8(0.19)© 3.6(0.24)® 3.9(0.17)4
A. falcata(2x) 12 4.6(0.19)8 2.6(0.14) 1.8(0.10) 32.0(0.48)8 3.0(0.00)* 1.0(0.00)° 2000.27)"
A. gardneri(2x) v 3.1(0.14)° 255 O21)e 13 (Oly Be 19.7(0.45)8 17 (O119) 2 4.64(0.46)8 1.4(0.19)8
A. gardneri(4x) 10 AUIN (OP cag 5-0(03 tyro Pe (O:1.2) Pe 45 el o2 1.5(0.05)° 3.8(0.12)8 1.9(0.37)8
A. tridentata(6x) 13 4.6(0.12)8 3.9(0.16)°© E201): 72100:39)" 2 OO 15 )e 8.5(0.45)* 2.0(0.49)®
* Volume = length x (width).
> Prominence of central apical tooth was scored 0 to 3 with central tooth absent = 0, = 1, conspicuous = 2, prominent = 3.

© Number of lateral appendages was scored 0 to 5 with no appendages = 0, numerous a caer =
PP & PP &E app

SdATv[aI Jvau sit pue iy]yeIINU xXaTdi7y ‘NOSHAANVS GNV ZLALS

10

202 Sipa 1L8(1)

Fic. 4. Fruiting bracts of F Atriplex vane and its near latves, A, A. nuttallii. B, A. See
°, A. cuneata. D, A. gardneri. E, A. falcata. Although there is considerable variation in the

ea)

fruiting bri acts of plants within and oe ips those shown are fairly represen-
tative of each species. Collection sites for those shown are: A. nuttallii, as W of Battle
Mountain, Lander Co.: Nevada; A. tridentata, Rush Valley, Tooele Co.: Utah; A. cuneata,
Navajo Mine lease site, ca 1O km SW of Farmington, San Juan Co.: New A. gardnert,
ca 10 km S of Bridger, Uinta Co.: Wyoming; A. falcata, ca 5S km N of Ontario, Malheur

Co.: Oregon. Bar = 15 mm

STUTZ AND SANDERSON, Atriplex nuttallii and its near relatives 203
RESULTS AND DISCUSSION

Probably because they are dioecious and wind-pollinated, most of the
shrubby species of Atriplex in North America are unusually rich genetically
and can often be best defined by quantitative, in addition to available sega
attributes. This is particularly true for A. nuttallii and its relatives. As show
in Figures 3, 5—8 and Tables 1—4, although A. nattallii and each of its near
relatives can usually be distinguished from each other by some qualitative
differences, when combined with differences in quantitative attributes and
geographic distributions, they are quite distinct.

Atriplex nuttallii $. Watson, Proc. Amer. Acad. moe 9:116.1874. Lecro-
TYPE: NEVADA: Reese Valley, Jul 1868, Watson 981 (GH

Phenotypically, A. zvttallii appears to be most oe, related to A. tridentata.
Although highly variable in A. nattalli (Fig. 2), most fruiting bracts of
both taxa have multiple, terminal, marginal teeth. Both have oblong or
oblanceolate leaves and occupy similar habitats (primarily saline bottom-
lands). Both are hexaploids (2”7=54) (a few tetraploid populations of A. tridentata
have also been found). A. nzuttallii differs from A. tridentata in being taller
statured (30—60 cm vs 10-30 cm) (Table 1), woodier, and more phenotypi-
cally variable. Much of its variation appears to be the result of hybridiza-
tion and subsequent introgression from other species, including A. tridentata
with which it is often sympatric. Plants of A. tridentata do not produce sa-
ponins whereas some plants | of A. nuttallii do, some do not (Table 3). A.
tridentata usually s} t-sprouting whereas A. zttallii is distinctly
caespitose with numerous 10" 120), slender (1—3 mm in diameter), woody
stems emerging from a single woody crown, 10—50 cm in diameter. Geo-
graphically, A. tridentata is common in northern Utah, southwestern Wyo-
ming, northwestern Colorado and northeastern Nevada (Fig. 8); A. nattalli
appears to be restricted to the alkaline valleys in northern Nevada and north-
western Utah (Fig. 3

AGES cuneata A. Nelson, Bot. Gaz. 34:357.1902. Typz: UTAH. fee RY
).: Emery, alt. 7,000 ft, 1894, M.E. Jones 5443 (HoLorype: US!; sorypes: MO! N
Hah,
ve — Rydb., Bull. Torrey Bot. Club 31:403. 1904. Type: COLORADO:
3 Sep 1897, J.H. Cowen 4071 (HoLotyee: US!; isotype: GH!).

Geoulsce®: i. cuneata are restricted primarily to eastern Utah, west-
ern Colorado and northwestern New Mexico (Fig. 5). In many places they
constitute the dominant vegetation on thousands of acres. The common name
of A. cuneata, “Castle-Valley clover,” connotes its value as a range plant, partly
because it is both palatable and nutritious for livestock and wildlife and
partly because, were it not there, there would apparently be nothing at all,

TABLE 3. Cytolo

fs

ined is in parenthesis.

gy, Saponin production,

flavonoid production, and geographic distribution of Atriplex nuttallii and its near relatives. The number of plants exam-

Taxon 2n Saponins Flavonols Geographic Distribution Date of
Chrom. # S 6-MeO 3-MeO Anthesis

A. nuttallii 54 (11) +,- (11) . - (21) valleys in n NV and nw UT Jun—Jul
A. cunedta 36 (126) - (69) + - (43) e UT, w CO, nw NM Apr-May
A. cunedta 18(87) -(72) - +(62) e UT, w CO, nw NM Apr—May
A. falcata 18 (115) + (74) - +,- (66) sID,n UT, sw WY, n NV, s OR, nw CA May—Jun
A. gardneri 18 (183) += (32) 7 + G1) MT, WY, AB, SK May—Jun
A. gardneri 36 (250) +,- (63) + - (40) MT, WY, AB, SK May—Jun
A. tridentata 54 (257) - (111) + - (105) n UT, n NV,s WY,s ID, nw CO Jul-Aug

bOC

(DST vals

STUTZ AND SANDERSON, Atriplex nuttallii and its near relatives 205

TaBLe 4. Key to Atriplex nuttallii and its near relatives [Series Nuttallianae of Standley (1916) con-
sisting on ee in which the plants are low-statured (less than 6 dm), suffrutescent ae nial, dio-

ecious, shrubs. Leaves are densely furfuraceous, alternate, entire, with Kranz-type anatomy. Fruiting
bracts are pres mm long, 2—5 mm broad, usually longer than broad and usually appendaged. }

1. Leaves linear, length more than 5 times the width
2. Root-sprouting extensive—A. tridentata
2. Root- ee none or minimal
Ss

3. Stems 50 or more from a woody crown, each 30—80 cm long—A. nuttallii
os ms single or few, each less than 30 cm long—A. falcata
l. tees ovate to oblong, length less than 5 times the width

2. Fruiting cae globose, 3-5 mm long with numerous flattened appendages—A. cuneata
2. Fruiting bracts elliptical to ovoid, 1-3 mm long with few appendages—A. gardneri

of forage value. This is conspicuous in many places where populations of A.
cuneata terminate abruptly against clay slopes that are completely devoid of
vegetation. It is also evident in sites where populations of A. cuneata have
experienced large-scale “die-back” and remain empty until repopulated by
new A. cuneata plants.

Atriplex cuneata is mostly tetraploid but throughout its range there are
several morphologically distinct diploid populations (Fig. 5). The variation
present in disjunct populations of tetraploid A. cuneata is probably the result
of introgression from these diploids, or in some cases, tetraploid A. cuneata
may have originated polyphyletically from different diploid ancestors.

Atriplex cuneata plants are mostly caespitose and erect with no evidence
of root-sprouting or layering. Their fruiting-bracts are usually much larger
than the fruiting bracts of A. gardneri (Table 2) and are covered with nu-
merous lateral appendages (Table 2, Fig. 4). They are usually free of saponins,
with exceptions only in populations in which introgressive hybridization
from other species is suspected. Tetraploid A. cuneata plants test positively
for the presence of 6-methoxy flavonols and negatively for the presence of
3-methoxy flavonals. In contrast, diploid A. cuneata plants test negatively
for the presence of 6-methoxy flavonols and positively for the presence of
3-methoxy flavonols (Table 3).

a a care (M.E. Jones) Standley, N. Amer. Fl. 21:68. 1916. Tver:
DAH ASHINGTON Co.: Weiser, alt. 2,000 ft, 7 Jul 1899, M.E. Jones 5.2. (HOLO-
TYPE: a ISOTYPE: UC!).

Populations of A. falcata are sporadically common in southern Oregon,
northeastern California, northern Nevada, northern Utah, southwestern
Wyoming and southern Idaho (Fig. 6). It is mostly diploid (27=18), but
occasional tetraploid populations occur (Pope 1976).

Phenotypically, A. falcata is most easily recognized by its small stature
(ca. 13 cm tall X 25 cm broad), small, linear leaves (ca. 24 mm long x 4
mm wide), and small fruiting bracts (ca. 4.5 mm long x 2.5 mm wide),

206 SIDA 18(1)

a
1 |
UTAH
i 1
' i
|
I fee eee eo ce - een= = at ama
eae
e ”
| Qa ey Lesa
| () )
e :
a)
| ny eo
! Shape © |
' ) '
“ ® e |
i \ QO : |
| . |
! \
{
e é COLORADO |
: pants dies Pet oo ee nannies net
O A. cuneata 2x ‘fo NEW MEXICO
| r ~
1 \ '
@ A. cuneata 4x C ee
—
oe See ae ee Sete ee ne Sone DA eee
Fic. 5. Geographic distribution of chromosome counts of plants of diploi oid (27=18) and

tetraploid (27=36) Atriplex cuneata.

usually without appendages, and terminating in a distinct, acute, central
apex (Tables 1, 2, Fig. 4)

The leaves of A. falcata produce abundant saponins (Table 3), a trait that
can be useful in field identification either by blowing bubbles in a water
emulsion of leaves or by tasting a leaf as it is chewed. When saponins are
present, bubbles form readily in a leaf-emulsion and the leaves have a dis-
tinctly bitter taste. Both of these tests are positive for A. falcata leaves, negative
for leaves of A. tridentata plants with which A. falcata plants are sometimes
confused. A. falcata plants are also readily distinguished from A. tridentata
plants by their rooting habit: A. tridentata plants are vigorous root-sprouters,
whereas A. falcata plants are caespitose with a single prominent taproot.
Also, A. falcata plants produce flowers and fruits early in the spring (May—
June) wheteas A. tridentata plants flower and set fruit mostly in mid to late
summer (July—August) (Table 3).

Atriplex gardneri (Mog.) Standley, N. Amer. Fl. 21:66. 1916. Tyr: {State?}.
La Platte River [date?]}, Gordon 250 (uoLotyPe: GH!, fragments).

Obtone gardneri Mog. in DC, Prod. 13:114. 1849.

Populations of A. gardneri (2x, 4x) dominate thousands of acres of clay
slopes and swales throughout much of southern and eastern Wyoming, central
and eastern Montana, western North and South Dakota, southern Alberta,

SruTZ AND SANDERSON, Atriplex nuttallii and its near relatives 207

e
ce]
e
eo.
of. hae
e ry -
®
e ‘
e e
Cd e
gece : ou e
_@ 9 °
°° e e° °
e rae e
e
e e @ce
e A. falcata 2x
e
(| A. falcata 4x

Fic. 6. Geographic distribution of chromosome counts of plants of diploid (27=18) and
tetraploid (27=36) Atriplex falcata.

and southern Saskatchewan (Fig. 7). Most populations are tetraploid but
diploid ones are common. In southern Alberta, southern Saskatchewan and
north-central Montana, diploid plants can usually be distinguished from
tetraploid plants by their smaller stature, smaller, thinner leaves, finer-tex-
tured, less woody stems and smaller fruiting bracts. However, throughout
most of the saline deserts of Wyoming and southern Montana, diploid and
tetraploid plants are phenotypically very similar although both show con-
siderable phenotypic variation both within and between populations. Chro-
mosome counts of plants in populations bordering Interstate Highway 80
in southern Wyoming, showed a preponderance of tetraploids west of the
Continental Divide and a preponderance of diploids to the east (Fig. 9).
However, no conspicuous ecological differences were evident between the
sites occupied by tetraploids and diploids, nor was it possible to consis-
tently distinguish diploids from tetraploids, phenotypically. This was true
for plants in natural populations as well as garden-grown specimens. They
are, however, clearly distinguishable by flavonoid differences: tetraploids

Stipa 18(1)

208
O
O
O
Ss)
© oO
@o
©
e
Qa
eo) O
oO oO
i. all |
os ° o8 |
em
%e O 0
6) OO O
\ e ° |
is e
+ e
®
e)
e {
e e ° °
e
°°
“~ e
’ oe
* Saebeheuci § Fy
o A.gardneri2x oe.
. e
e A. gardneri 4x
Fic. 7. Geographic distribution of chromosome counts of plants of diploid (27=18) and

tetraploid (27=36) Atriplex gardnert.

always produce 6-methoxy flavonols, diploids do not; diploids produce 3-
methoxy flavonols, tetraploids do not (Table 3). Although both diploids
and tetraploids sporadically produce saponins (Table 3), diploids are most
often heavy producers, tetraploids usually produce little or none.

Atriplex gardneri plants are short-statured, mostly 4-10 cm in height,
(Table 1), but are often more than 80 cm in diameter. The plants are cae-
spitose, arising from a single deep taproot but often show considerable lay-
ering. Their fruiting-bracts are usually smaller (ca. 3—4 mm) and have fewer
lateral appendages than those of most near relatives (Table 2, Fig. 4).

STUTZ AND SANDERSON, Atriplex nuttallii and its near relatives 209

\ r 1
, ! 1
/ he \
if ae \
us mM Wl.) eee eee Oeil eee ae 4
1 “ |
a ae \! |
i i} ‘
/ i)
i
i | r
H | 1
I i PS anu oceans 7
i ! 1
' ! 1
' { \
! | !
| \
Nata pate re I e \ e e@ \
eo. ne -
| aaa E p----~@----- @---| @ |
sek go
pe Le @ @e !
1 ! H e
| | 8 e ‘ @ A eee :
' } (ll @-- eee eee \
' ! e \
H e %Y eMme e
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! I #3 ( H e b=
i \
t e ' e ga 1 1
‘ | e e ! 1
2 | |
r
' : \
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| !
i @ \
{ |
7 | \
N \
: (5-7 coe | nena nner pegcat es
* A. tridentata 4x } aaa
sella etic ema Dane |
! \
4 } \ non
e@ A. tridentata 6x is | | r
_ ie. \ |

8. Geographic distribution of chromosome counts of plants of tetraploid (27= 36) and
eee (2n=54) Atriplex tridentata.

pele tridentata Kuntze, Revis. Gen. a 2:546, 1891. Tyre: UTAH: Corinne,
_E.O. Kuntze 3084 (HOLOTYPE: NY

Two nee races of Atriplex oo have been found: tetraploid
(2n=36) and hexaploid (27=54). Hexaploids are, by far, the most common,
occurring throughout much of western United States and in most of the
valley bottoms in Utah and Nevada that were recently occupied by Pleis-
tocene lakes (Fig.8). Only three major tetraploid populations are known:
one in Juab County, Utah, one in Eureka County, Nevada, and one in Lin-
coln County, southwestern Wyoming (Fig. 8). Although tetraploid and hexaploid
A. tridentata are to some extent, morphologically distinct, the differences
do not appear to be sufficient to warrant designation as separate taxa.

Atriplex tridentata differs from other Atriplex species in several features,
including linear to oblong, furfuraceous leaves, fruiting bracts with con-
spicuous terminal, marginal teeth (Fig. 4), late-flowering habit July-Au-
gust), and vigorous root-sprouting. A. tridentata plants are mostly herba-
ceous above ground with a few slender stems growing erect from woody
underground crowns and roots.

The extent of root-sprouting in A. tridentata plants is often made con-
spicuous by its dioecious flowering habit. By noting its flowers, the extent
of a single male or single female plant may be easily determined. In some
cases individual plants have been found to occupy more than 200 m~. Sometimes

210 Sipa 181)

Little Green
America River

je ny | r ; . aE
90 95 LOO 105 110 115 120 125 130 135

t
Rock Point of
Springs Rocks

Z oo ffllaballes

I Sree Os
150 155 160 165 170 175
Wanisutter
| eur —mlles flea ie tle le
180 185 190 195 200 t 205 210 215 220 225
Rawlins

SLED aa ym,
225 230 235 240 245 250 255 260 265 270
. 9. Distribution of diploid and tetraploid populations of Asriplex gardneri alongside
nan highway 80 in souther oming from the Utah border (mile 0) to near Elk
Mountain Wyoming. C eres counts were made on plants at each mile marker, when

available, seapled . = diploids, black bars = tetraploids.

a single plant forms a complete carpet in which there are no other plants; at
other times the root-sprouting carries a plant around and between plants of
other species.

Possibly because of its low saponin content (Table 3), its root-sprouting
habit and predominantly herbaceous tissues, A. tridentata is often regarded
favorably by ranchers as forage for livestock (personal reports).

STUTZ AND SANDERSON, Atriplex nuttallii and its near relatives 211
ACKNOWLEDGMENTS

The authors thank Dr. Richard W. Spellenberg and Dr. Ronald L. Hartman
for numerous helpful suggestions, Broken Hill Proprietory Minerals and
Brigham Young University for financial assistance and the curators of the
following herbaria for loans of specimens and access to their collections:

BRY, CAS, GH, MO, NY, PH, RM, RSA, UC and US.

REFERENCES

Bassett, LJ.,C.W. Crompton, J. McNett, and P.M. TascHEREAU. 1983. The genus A¢riplex
(Chenopodiaceae) in Canada. Monograph No. 31. Communications Branch, Agriculture
Canada, Ottawa

Greuter, W., H.M. Burpet, W.G. CHALONER, V. DEMONLIN, R. GROLLE, D.L. HAWKsworTH,
D.H. NicHotson, P.C. Sirva, F.A. Starteu, E.G. Voss, and J. McNer. 1988. Interna-
tional code of botanical nomenclature. Adopted by the ail International Botani-
cal Congress, ae eee oa oae eae Regnum Ve

ee E, H.M.B T, W.G. CHALONER, 7 “s MOULIN, D.L. HawKsworTH,

P.M. JORGENSEN, D. 7" Soe "PC. Sitva, P. TREHANE, and J. McNem. 1994. I
national code of botanical nomenclature (Tokyo C ode). Adopted by ne Scat ae
national Botanical Congress, pee August-September 1993. Regnum Veg. 1

Hai, H.M. and FE. Crements. 1923. T. ee ae method in taxonomy. Carnegie
has of Washington Publication No. 326.

Hanson, C.A. 1962. Perennial Atriplex of Utah and the northern deserts. Thesis. Brigham
Young University, Provo, Ut

McNEIL, J., I,J. Basserr, C.W. Crompton, and P.M. . 1983. Taxonomic and nomenclatural
notes on Atriplex L. (Giese Taxon 32: 549-556.

Nutta.l, T. 1818. The genera of North American plants. Vol. 1. Philadelphia. D. Heartt.

Popr, C.L. 1976. A phylogenetic study of the 2 aa shrubs in the genus Afriplex.

Ph.D. Dissertation, Brigham Young University, Provo,

Pursu, E.T. 1814. Florae Americae Sepicaeenle Vol. I. ee White, Cocchrane and
Company

SANDERSON, S.C., and H.C. Stutz. 1984. Flavonoid aglycones of diploid and saat ed
confertifolia. In: A.R. Tiedemann, E.D. McArthur, H.C. Stutz, R. Stevens L. Joh
K.L. (compilers). Proc. symposium on the biology of Atriplex and ae ee,
Provo, UT.

, R.L. PeNpLETON, E.D. Mcarruur, and K.T. Harper. ae a effect
ipl

iS]
es

n,

on a seal mammal forage preference ina payee of At F.J. Provenza,
J.T. Flinders and E.D. McArthur (compilers), P eropoe tia on See Herbivore

Interactions. USDA Forest Service General Technical Report INT-222, 74-77. Ogden,

STANDLEY, P.C. 1916. Chenopodiales. North American Flora 21(1). New York. New York
Botanical Garden

Stutz, H.C. 1978. pentose evolution of perennial Atriplex in western America. In
Harper fdlh I. Reveal, organizers. Intermountain biogeography: A symposium. Great
Basin Naturalist Mem.. 2:161-168.

ee ,C.L. Pope, and S.C. SANDERSON. 1979. Evolutionary lies of Atriplex: Adaptive
products from the natural hybrid 6N A. tridentata __4N A. canescens. Amer. J. Bot. 66:1181—
1193

212 Sipa 18(1)

Urpricu, E. 1934. Chenopodiaceae. In: A. Engler and K. Prantl, eds. Die Natiirlichen
Pflanzenfamilien. Ed. 2. 16C:379-584. Engelmann, Leipzi
WatTSON, S. 1871. Botany. In: C. King. Report of geological apleaies of the fortieth
parallel. Vol ume V. U.S. Government Printing Press, Washington, D.C.
. 1874. A revision of the North American C
Pese. oR 82-1 26.
WesH, S.L. 1984. Utah flora: Chenopodiaceae. Great Basin Naturalist 44:183—209.

—

nenopodiaceae. Proc. Amer. Acad.

FLAVONOIDS IN STROPHOSTYLES SPECIES
AND THE RELATED GENUS DOLICHOPSIS
(PHASEOLINAE, FABACEAE):
DISTRIBUTION AND PHYLOGENETIC
SIGNIFICANCE

JUAN PABLO PELOTTO and MARIA A. DEL PERO MARTINEZ

Centro de Estudios Farmacolégicos y Botdnicos
Consejo Nacional de Investigaciones Cientificas y Técnicas
Serrano 665, Buenos Aires, 1414, ARGENTINA

ABSTRACT

Strophostyles Elliott is the only genus within the Phaseolinae with a center of distribu-
tion in the United States. It comprises three species, namely S. he/vwla (L.) Elliott, 5. vnbellata
(Willd.) Britton, and S. lerosperma (Torrey & A. Gray) Piper, and it is considered as allied to
Dolichopsis Hassler, a monotypic genus endemic to South America. This study analyses the
leaf flavonoid content from the three es ppecic™ and akin es paraguariensis on
ee De aim mC RNAS the payige 2 g taxa, We isolated 38 gly-

I in. All St phostyles species were characterized

by the presence of isoiaamimedia glycosides. However, S. Diowere showed a distinctive
profile while 8. helvwla and S. umbellata clustered together. In contrast, D. paraguartensis
lacked isorhamnetin-based compounds. A cladistic analysis of flavonoid plus morphologi-
cal data supported ak monophyly and showed S. /eiosperma as the sister taxon of
the clade S. helvula-S. umbellata.

RESUMEN

Strophostyles Elliott es el nico género de las Phaseolinae con un centro de distribucién
en los Estados Unidos. Comprende tres especies: S. helvula (L.) Elliott, S. umbellata Willd.)
Britton y S. be alg ney . ae Cry) Piper, y se considera afin a Dolichopsis Hassler, un
género Este estudio analiza el contenido de flavonoides

foliares « en ieee tres especies = Strophostyles yen Bees paraguariensis Hassler con el objeto
Se aislaron 38 glicésidos de ae
| racterizaron por la presencia

de examinar |

quercetina e isoramnetina. Todas
de ey de isoramnetina. Sin caer Ay eioperna mostré un perfil ee mientras
eS. hele wla and S. umbellata se agruparon juntas. Por el contrario, D. paraguariensis no
basados en la isoramnetina. Un andlisis cladistico conjunto de los dat
de Aavanoides y caracteres oie apoy6 la monofilia del género y mostr6 a8. lesosperma
como el tax6n hermano del clado S. helvula-S. umbellata.

a [Pan

INTRODUCTION

Strophostyles Elliott is the only genus within the Phaseolinae with a cen-
ter of distribution in the United States. Its current taxonomic treatment
follows the original generic concept (Elliott 1822), but it was previously

Sipa 18(1): 213-222. 1998

214 Sipa 18(1)

associated with unrelated species and reduced to a section of Phaseolus (de
Candolle 1825; Bentham 1837, 1865) until it was restored as a separate
genus including three species (Britton & Brown 1897; Piper 1926). A set
of characters precludes merging Strophostyles with Phaseolus, namely erect
style (not coiled), lack of hooked hairs, pedicels shorter than the calyx, and
nodes of the inflorescence somewhat swollen (Maréchal et al. 1978). More-
over, Strophostyles plants can be recognized by their nearly asymmetric flowers
arranged in subumbellate inflorescences, bracts and bracteoles persisting
through seed maturation, cylindrical seeds (often pubescent), and linear pods.
Strophostyles has also been considered as allied to Dolichopsis Hassler (Maréchal
etal. 1978; Lackey 1983). Nevertheless, both genera are easily distinguishable
by many characters and their quite distinct geographical distributions. The
monotypic genus Dolichopsis! resembles Strophostyles mainly in floral mor-
phology (purplish corolla, keel with a not curved to somewhat curved beak,
style slightly thickened distally, and stigma terminal oblique) and general
appearance, but it is unlike the latter in having symmetric flowers clustered
in elongate pseudoracemes and the unique fruit traits such as oblong, very
flat pods with oblong seeds implanted through a very long funicle and with the
hilum perpendicular to the placenta. The geographic range of Dolichopsis is Paraguay
and Argentina, in South America, while Strophostyles occurs throughout eastern
USA, eastern Canada up to south of Quebec and extreme northeastern Mexico.
Phytochemical data on Strophostyles species are scanty and include the absence
of both leuco-anthocyanins (Baudet 1978) and canavanine (Lackey 1977),
and a recent report of flavonoids (Williams et al. 1995). In this study, we
expanded on the survey of foliar flavonoids by considering a larger number
of samples of the three Strophostyles species, i.e. S. heluula, §. umbellata and S.
lerosperma, and we added Dolichopsis paraguariensis for comparison.

MATERIALS AND METHODS

We analyzed the constitutive flavonoids present in the leaves of herbarium
specimens belonging to the three Strophostyles species and Dolichopsis paraguariensis.
Samples (100-200 mg) were powdered and extracted under reflux with 80%
methanol (x 3). Concentrated methanolic extracts were two-dimensionally
chromatographed on paper (BAW/ 15% acetic acid). Compounds were identified
by standard methods (Mabry et al. 1970; Markham 1982). These included
complete and controlled (3 min.) acid hydrolysis, enzymatic hydrolysis (-
glucosidase), co-chromatography with authentic markers and UV-Vis spec-

'Dolichopsis was a genus with two species, D. paraguariensis and D, monticola (Lackey 1983,
Lewis 1991), but recently Delgado Salinas & Lewis (1997) created the new genus Oryxis
where they placed D. monticola. Therefore, D. paraguariensis has became the unique repre-
sentative of the genus.

PELOTTO AND Martinez, Flavonoids in Strophostyles and Dolichopsis species 215

troscopy. Glucosides were separated from their galactosidic analogues by
TLC in the appropriate system according to Budzianowski (1991).

Plant material.—Specimens were provided by the Instituto de Botanica
Darwinion Herbarium (SI), San Isidro, and the Centro de Estudios Farmacol6gicos
y Bordnicos Herbarium (BACP), Buenos Aires.

Strophostyles helvula (L.) Elliott

U.S.A. Arkansas. Jefferson Co.: Arkansas river bottoms, 220 ft, 17 Sep 1937, Demaree
16245 (SI). Ilinois. Mc Donough Co.: Argykle Lake, near Colchester, 2 Aug 1958, Jones
22335 (SI). lowa. Dickinson Co.: N shore of Spirit Lake, sand (older beach), 5 Aug 1913,
Sinek 14... (number illegible) (SI. Mississippi. Harrison Co.: near the coast on sand, 6
Jan 1951, Demaree 30675 (SI); Ship Island, P.O. Biloxi, in stabilized sand, moist, long trailing,
Demaree 31059 (SI). Virginia. Prince George Co.: rich alluvial thicket back of sand-beach
of James River, Jordan Point, SE Virginia, 16 Sep 1938, Fernald & Long 9353 (SI). Un-
known locality:{Flora of the Western Reserve} 15 Aug 1897, G.B. Ascheroft, Berea O. s.n. (SI).

Strophostyles leiosperma (Torrey & A. Gray) Piper

U.S.A. Oklahoma: 5 mi NW of Breckenridge, 25 Jul 1941, Gephardt 747 (SI). Woods
Co.: in waste place, hard soil, near Alva, 24 Sep 1913, Stevens 2824 (SI). Texas. Smith Co.:
Amigo, neglected sandy field, 10-17 Aug 1945, Moore, Jr. 995 (SI). Morris Co.: Aug 1891,
Carleton 420 (SD.

Strophostyles umbellata (Willd.) Britton
U.S.A. Virginia. Greensville Co.: dry pine and oak woods, about | mi N of Skipper’s,
14-15 Jul 1938, Fernald & Long 8737 (SI).

Dolichopsis paraguariensis Hassler

ARGENTINA. Entre Rios: Depto. La Paz, R 126, desvioa Ombtes, borde camino, 31
Jan 1981, fl. azul-violdceo, Troncoso de Burkart & Bacigalupo 3096 (SI). PARAGUAY. Depto.
Pte. Hayes: Estancia Loma Pyta, 23° 40'S, 59° 35'W, 2 Apr 1974, enredadera casi rastrera,
fl. violéceas, crece en pastizal, n.v. ‘kekleichetas,’ Arenas 544 (BACP). Depto. Boqueroén:
Misién Santa Rosa, 21° 45'S, 61° 35'W, Feb 1981, enredadera, fl. violaceas, crece en pajonal,
n.v. ‘ceihlowéy’, Arenas 1726 (BACP).

Data analysis.—A cluster analysis was performed on flavonoid data of
15 herbarium specimens. Similarity matrix was measured using Jaccard’s
coefficient and a dendrogram was constructed applying the unweigthed pair-
group method of arithmetic averages (UPGMA). All calculations were done
using NT-SYS program (Rohlf 1993).

A cladistic analysis of the flavonoid data plus a set of morphological characters
was carried out according to the maximum parsimony principle. Outgroup
criterion was used for character polarization. Vigna adenantha was chosen as
the external group which flavonoid data were obtained following the methods
above mentioned (Pelotto, unpublished manuscript). For flavonoids, char-
acter states that occurred in the outgroup were scored as 0 and those in the
ingroup (D. paraguariensis and the three Strophostyles species) were scored as
1 (see Appendix, Tables A and B). Morphological data were gathered from
the literature and included some multistate characters that were treated as
non-additive (see Appendix, Tables A and C). Cladograms were calculated

216 Stipa 18(1)

using the implicit enumeration routine (i.e.*) of the program Hennig86
(Farris 1988) wich all characters equally weighted.

RESULTS AND DISCUSSION

Chromatographic properties of the identified flavonoid glycosides and
its distribution in Strophostyles species and D. paraguariensis are shown in
Tables 1 and 2, respectively. All detected compounds were O-glycosides of
flavonols with sugars attached at positions 3 and 7 of the aglicone skeleton.
This pattern of glycosilation is very common among the Phaseolinae (Zallocchi
& Pomilio 1994; Williams et al. 1995, Pelotto unpublished manuscript).

All three Strophostyles species produced glycosides based on the methy-
lated flavonol isorhamnetin plus kaempferol and quercetin. Notwithstanding
S. helvula and S. umbellata showed very similar chromatographic patterns,
while the flavonoid profile of S. /e/osperma was quite distinctive. No rhamnosides
were detected in S, /esosperma and it only shared the presence of monoglycosides
with the other two species. Based on a three-sample analysis within a fla-
vonoid survey of the Phaseolinae, Williams et. al (1995) have also reported
the occurrence of isorhamnetin glycosides in Strophostyles species and no-
ticed the same interspecific differences. In contrast with our results, Will-
iams and co-workers isolated fewer compounds and did not detect kaempferol
glycosides from leaves, although they did from stems and/or flowers.

In turn, Dolichopsis paragnariensis samples were characterized by the presence
of kaempferol and quercetin glycosides, lacking isorhamnetin, Noticeably,
Paraguayan samples contained only kaempferol glycosides while Argentinean
one had kaempferol plus quercetin glycosides. However, ina previous work
(Zallocchi et al. 1995) both kaempferol and quercetin glycosides were re-
ported from one sample of D. paraguariensis from Paraguay, but of the eight
flavonol glycosides the authors identified only rutin and kaempferol-3-O-
rutinoside were also present in our samples. These differences may be due
to the fact that Zallocchi and co-workers analyzed a whole plant extract
and therefore their results are difficule co compare with ours.

After the cluster analysis S. helvula and S. umbellata are closer to D. paraguariensis
than to S. /esosperma (Fi g. 1). This is because S. helvala and S. umbellata have
more glycosides (based on kaempferol and quercetin) in common with D.
paraguariensis than with S. le‘osperma, even though D. paraguariensis does not
produce isorhamnetin glycosides.

Cladistic analysis resulted in two most parsimonious trees (length, L =
35, consistency index, CI =91, retention index, RI =70. Fig. 2). Both cla-
dograms support Srrophostyles monophyly but differ in the depicted rela-
tionships among Strophostyles species. One tree (Fig. 2.A) shows S. helvula
and S. /esosperma as being sibling species, but this hypothesis needs the par-
allel gain of the characters 19, 22 and 31 on the S. wmbellata and S. helvula
branches. The other tree (Fig. 2.B) supports the clade 8. umbellata-S. helunla

PELOTTO AND Martinez, Flavonoids in Strophostyles and Dolichopsis species 217

TaBLe |. Chromatographic characteristics of the identified compounds.

COLOUR* Rf (x 100)
SPOT IDENTITY UV + NH3 BAW 15% aa
1 K-3-O-glucoside + K-3-O-galactoside DP G 69 45
2 K-7-O-glucoside + K-7-O-galactoside Y Y 45 15
3 K-3-O-rutinoside + K-3-O-robinobioside DP Y 52 54
4 K-3-O-diglucoside + K-3-O-digalactoside DP Y 33.5 61.5
5 K-3,7-O-diglucoside DP Y 29 70
6 K-3-O-rutinoside-7-O-glucoside +
K-3-O-robinobioside-7-O-glucoside DP Y. 24 75
7 K-3,7-O-triglucoside DP Y 3, 82
8 K-3,7-O-triglycoside (glu + rha + gal) DP ¥ 4 80
9 Q-3-O-glucoside + Q-3-O-galactoside DP ng 56 39
10 Q-7-O-glucoside + Q-7-O-galactoside Y YO Dif 9
11 Q-3-O-rutinoside + Q-3-O-robinobioside DP Y 40 51
12 Q-3-O-diglucoside + Q-3-O-digalactoside DP Y 26 52
13. Q-3,7-O-diglucoside DP ¥ 23 63
14. Q-3-O-rutinoside-7-O-glucoside +
Q-3-O-robinobioside-7-O-glucoside DP Y 15 70
15 Q-3,7-O-triglucoside DP Y 3 78
16 Q-3,7-O-triglycoside (glu + rha + gal) DP Yo 5) 79
17 IR-3-O-glucoside + IR-3-O-galactoside DP x 56 42
18 IR-7-O-glucoside + IR-7-O-gal si Y ¥ 38 10
19 IR-3-O-rutinoside + IR-3-O-robinobioside DP ¥ 40 54
20 id 3- O-diglucoside + IR-3 -O-digalactoside DP Y 29 58
21 3,7-O-diglucoside DP Y 2) 68
22 oa de-7-O-glucoside +
IR-3-O-robinobioside-7-O-glucoside DP Y 19 74
23 IR-3,7-O-triglucoside DP 3 80

"DP: deep purple, Y: yellow, YO: yellow- uae:
‘glu: glucose, rha: rhamnose, gal: galactose

and requires three reversions (characters 1, 9 and 17). This scenario is pref-
erable to that portrayed on Fig. 2.A since a mutation lost is a more prob-
able event than the homoplastic acquisition of isorhamnetin glycosides. Even
more, if we suppose reversal of characters 1, 9 and 17 on the S. wmbellata
branch as being a consequence of sampling error (undersampling), the cla-
dogram becomes shorter with only 32 steps (CI= 100, RI= 100) and the
unique solution of a similar analysis. Thus, we consider the tree depicted
on Figure 2.B a more plausible ingroup phylogen

Flavonoid evolution shows methylation of the davousl skeleton as an advanced
character shared by all Strophostyles species and the absence of rhamnosides
in S. lezosperma as an (aut)apomorphic loss.

Morphological traits are congruent with flavonoid data. Subumbellate
inflorescence (character 24), persistent bracts and bracteoles (character 26),
linear, terete pods (character 27) and seed pubescence (character 28) sup-
port Strophostyles monophyly, and are correlated with isorhamnetin monoglycoside
production (characters 17 and 18). Strophostyles helvula strongly resembles

E 2. Glycoside distribution in the analyzed samples. Compounds are numbered according to Table 1. K: kaempferol glycosides; Q: quercetin glycosides; IR:

a. glycosides; +: present; -: absent
K IR
l 2 3 4 > 6 8 9 10 11 12 13 14 ii: 16 17 18 19 20. 21 22° 23

S. helvula
Jones 22335 - + . - + - + - - - + - - - - . - -
Sind 14. + + : - + = i Fs : : E a + + + - - 4 -
Aschroft & ames s.n. + + = = + 7 + + + - 7 + é + + + + - z i :
Demaree 30675 { if = - + = + + 2 7 + . + + + 2 - + =

emaree ie 5 - - - + - : 7 + = + 7 + - é + 7
Demaree 3 + + = ~ + - + + + - : + = + + ' t - : + 7
Fernald & ae 9353 . + - 7 + 7 - + + = = + - + 7 + ' = be + =
S. umbellata
Fernald & Long 8737 ‘ + + 7 - + 7 - + + z - + S + - + + : - + :
S. leiosperma
Moore Jr. 995 . - - + : 2 + + - : + : < + + 7 : a = zs
Carleton 420 - + + - - + = : + . + a + - : + + : ri
Gephardt 747 + - + + - - + - - + + - i 7 + 2 z + 4: E +
Stevens 2824 + - + + - - + 7 - + + : + 7 4 2 . + # < 4:
D. paraguariensis
Arenas 1726 - + z = + 2 y s 2 - : a zs e - - 7 : Z . =
Arenas 544 za 2 7 + 2 2 “ . : < - 7 - - . : z 7 _ 7
Troncoso & Bacigalupo 3096 - + 7 - + + - + + 7 - + é + 2 = 2 - _ 2 =

(181 ¥als

PELOTTO AND Martinez, Flavonoids in Strophostyles and Dolichopsis species 219.

JACCARD’S COEFFICIENT

0.00 0.25 0.50 0.75 1.00
HEL-22335

LEI-747
|

L

Fic. 1. Dendrogram of the Strophostyles and Dolichopsis specimens constructed from a simi-
larity matrix ernie s coefficient) using the UPGMA method. Cophenetic correlation co-
efficient, r = 0.96

S. umbellata, except for its more lobed leaflets and larger pods and seeds.
Strophostyles leiosperma is rather different from the other two species because
of the smaller flowers (character 25) arranged in more pauciflorous inflo-
rescences and its seeds glabrous and shining at maturity (character 29).

Similarly, both S. e/vula and S. umbellata are more widespread and northerly
distributed, with the first species reaching Canada, while S. /e‘osperma has a
more limited distribution ranging from south of the United States to the
extreme northeastern of Mexico (Britton & Brown 1897, Maréchal et al.
1978). Species divergence at chemical and morphological level also correlates
with their ecological features; S. helvula and S. umbellata mostly grow in
more mesic sites, while S. /ezosperma is adapted to live into more xeric habitats.

Regarding the evolution of the growth form, overlapping this character
onto our preferred topology suggests that annual growth would have evolved
independently in both S. /ezosperma and S. heluula, whereas perennation would
be the plesiomorphic state shared by D. paraguariensis and S. umbellata.

In summary, universal occurrence of isorhamnetin-based compounds in
Strophostyles species is a good chemical character in defining generic monophyly
while individual glycosides are useful characters to trace species evolution. With-
in the Phaseolinae isorhamnetin glycosides have sporadically been recorded
in four Phaseolus species (Pelotto, unpublished manuscript) and some Vigna
and Macroptilium species (Zallocchi & Pomilio 1994; Williams et al. 1995).
This fact suggests that flavonol methylation has appeared several times in
the tribe, making i it a valuable phylogenetic marker at the infrageneric level.

Beyond this ion, flavonoid data from the related genera Oxyrhynchus
and Oryxzs are wanting for a complete view of this little group of American
species around Vigna.

SIDA 18(1)

Ag %
$ F :
aS & SS &
S S x % u
= So 3 Ss &
% BS SS S S
NS S iS ‘3 3
& x S s =
S ~ oy vy “3
~ G Sse 223 “Sgi9za Sst
\ * Ne S — 1-15
* \ Sf “eet 23
~\ \ % > 25l2),29
\ we 8.27(1).30 a 9.17
7 ~< er 4
\ % a
o a wpe 18,24,26,27(2),28
Me af
a 4
. mote 2,10,29(1)
va
B-
wh
AY a
8 : :
: : : $
G SS S SS &
g No x 8 S
As) %S w S &
% S ‘S$ Sy S
S&S Q & 3 S
S S — ee MA
% fe 3 = ” Pi LA.
a X itel6 So 1917
. wee 20.2123 KY
‘ \ 28{2),29 Xe
. \ : |
sien 2°7(1),30 \ wpe 19.22.51
\ \ Se .
~*~ a Zs 18,24 26 27(2), 28
~~ — wo
~ a
mt 2,10.25(1)
13
Fic. 2. The two most parsimonious trees (L = 35) generated using the data matrix (see

Appendix, Table A) and Vigna adenantha as outgroup. Characters are mapped on the trees
as follows: solid bar = non-homoplasious apomorphy, clear bar = homoplasious apomorphy,
and cross = reversal. Numbers on the right of the character symbols stand for character
numbers (and character state).

PELOTTO AND MartTiNeEz, Flavonoids in Strophostyles and Dolichopsis species 22h
ACKNOWLEDGMENTS

We thank the curators of the Instituto de Botanica Darwinion Herbarium
(SI) and the Centro de Estudios Farmacolégicos y Botanicos Herbarium (BACP)
for providing the plant material, and CONICET for financial support. We
are also grateful to two anonymous reviewers for suggestions that improved
the earlier manuscript.

APPENDIXES

Taste A. Data matrix for the cladistic analysis including both flavonoid (characters 1-15, 17-23,
codified according to Table B) and morphological (characters 24—31, codified according to are eC)
data sets and using Vigna adenantha as outgroup.

character number

Taxon 12 3 4 5 6 7 8 9 1011 12 13 14 15 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31

Vigna adenantha — (

=
Oo
o
a
j=)
So
cS
co
So
o
Oo
—
oS
v
=
°

D. paraguartensts

S. helvula Tes 300 0:00.04 Ud

o
o
=)

S. umbellata

S. leiosperma [ies 81:2 al Aare 21 Sil “Oh ak’ Sie Mere lel gels oO ts sme) felasle ee2arpeliene2 1s 1 xO:1 0)

ABLE B. Flavonoids. Character numbers are as in Table 1. Character 16 was not included because no

hypothesis abour homology can be made on a partially identified compound.
Character states
1. O= absent, l= present 8. O= absent, l= present 15. O= absent, l= present
2. O= absent, 1= present 9. O= absent, 1= present 17. O= absent, 7 present
3. O= present, 1= absent 10. O= absent, 1= present 18. O= absent, 1= present
4. 0= absent, 1= present 11. O= present, 1= absent 19. O= absent, l= present
5. Q= absent, l= present 12. O= absent, 1= present 20. O= absent, l= present
6. is present, 1= absent 13. O= absent, 1= present 21. O= absent, 1= present
7. 0= absent, l= present 14. O= present, 1= absent 22. O= absent, 1= present
23. O= absent, l= present
Tasie C. Morphological characters, states and polarities.
character states
24. inflorescence 0= pseudoracemose; 1= subumbellate
25. flower O= great (> 20 mm); 1= medium (7-15 mm); 2= small (< 7 mm)
26. bract ea eer O= persisting no x anthesis; 1= persisting through seed maturation
27. pods = linear, compressed; 1= oblong, very flat, with false cellulosic septa; 2 =
linear, cylindrical
28. seed coat O= smooth; 1= pubescent
29. seed pubescence ?= naps O= persisting in mature seeds; 1= absent from mature
seeds

30. hilum O= parallel to che placenta; 1 = Ee ae to the placenta

31. leaflets O= entire; 1= lobed to somewhat lobed at

Sipa 18(1)

No
i)
No

REFERENCES

Baupbet, J.C. 1978. Prodrome d’une ee générique des Papilionaceae aseoleae.
ull. Jard. Bot. Belg. 48:183—
BENTHAM, G. 1837. Commentationes a leguminosarum generibus. J.P Sollinger, Vienna.

Bentham, G. 1865. Leguminosae. In: Bentham, G & Hooker, J.D. Genera Plantarum 1:434—
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00.

Britton, N.L. and A. Brown. 1897. An illustrated flora of the northern United States,
Canada and the British Possessions. Vol. II. Charles Scribner's Sons. New York

BuDZIANOWSKI, J. 1991. Separation of flavonoid glucosides from their galactosidic analogues

thin-layer Se aphy. J. Chromatography 540:469—474.

DE CANDOLLE, A.P. 1825. Prodomus systematics naturalis regni vegetabilis Vol. 2. Paris,
St about, oe

DELGADO SaLinas, A. and G.P. Lewis. 1997. Oryxis, a new genus in tribe Phaseoleae (Leguminosae:

Papilionoideae) from Brazil. Kew Bull. 52:221—

E.uorr, $. 1822. A sketch of the botany of South cane and Georgia. 2:229.

Farris, J.S. 198 88. Hennig86 version 1.5. Sic and software documentation. Published
by the author, Port Jefferson Station, New

Lackey, J.A. 1977. A revised classification of the cribe Phateciew (Leguminosae-Papilionoideae)
and its aera to canavanine distribution. J. Linn. Soc., Bot. 74:163-1

Lackey, J.A. 1983. A review of generic concepts in American Phaseolinae @abanes: Faboideae).
Iselya sear

Lewis, G.P. 1991. A new combination in Dolichopsis (Leguminosae: Papilionoideae). Kew
Bull. ghee

Masry, T.J., K.R. MARKHAM, and M.B. THomas. 1970. The systematic identification of
ones ene Verlag, New York

Maretcual, R., J.M. Mascuerpa, and F. Stainier. 1978. Etude taxonomique d’un groupe
complexe d’espéces des genres Phaseolus et Vigna (Papilionaceae) sur la base de données
morphologiques et polliniques, traiteés par l’analyse informatique. Boissiera 28:1—273

MarkuaM, K.R. 1982. Techniques of flavonoid identification. Biological Techniques Se-
ries. Academic Press. London.

Piper, C. V. 1926. Studies in American Phaseolineae. Contr. U.S. Natl. Herb. 22:663-701.

ROHL br, FJ. 1993. NTSYS-pc. Numerical taxonomy and multivariate analysis system, ver-

—

on 1.8. Exeter Software, Setauket, N-Y.

Witiams, C.A., J.C. ONYILAGHA, and J.B. HARBoRNE. 1995. Flavonoid profiles in leaves,
flowers and stems of forty-nine members of the Phaseolinae. Biochem. Syst. Ecol. 23:655—
667

ZALLOCCHI, E.M. and A.B. oe 1994. Evolution of flavonoids in the Phaseolinae. Phy-
tochemistry 37: 449-4

ZALLOCCHI, E.M., A.B. ois and R.A. eam 10s. 1995. Estudio oo de

a subtribu P Fgseolinie (Phaseoleae-Papilionoideae-Leguminosae) oe -

as especies argentinas de los géneros Piso: y Dolichopsis, Pari 33:135-

—

RUELLIA JIMULCENSIS (ACANTHACEAE),
A NEW SPECIES FROM THE CHIHUAHUAN
DESERT AREA, MEXICO

JOSE A. VILLARREAL Q.

Departamento de Botdnica
Universidad Autinoma Agraria “Antonio Narro”
Buenavista, Saltillo, Coahuila 25315, MEXICO

ABSTRACT
Ruellia jimulcensis Villarreal sp. nov., from the Jimulco mountains area (southwestern
Coahuila and northeast Durango), is described and illustrated. It is similar to R. occidentalis
(Gray) Tharp and Barkley, and recognized by its relatively smaller flowers, shorter corolla
basal tube and its distribution.

RESUMEN

Ruellia jimulcensis Villarreal sp. nov. del 4rea de las montafias de Jimulco (suroeste de
Coahuila y noreste de Durango), es descrita e ilustrada. Es similar a R. occidentalis (Gray)
Tharp y Bee y se reconoce por sus flores mas pequefias, tubo basal de la corola mas
corto y su distribucién.
Key Worps: Acanthaceae, Rve//ia, Coahuila, Flora of México.

Ruellia, a tropical and subtropical genus of about 200 species is repre-
sented in the Chihuahuan Desert Region by six species (Henrickson & Johnston,
in press). A new species is proposed as part of the study of the Flora of Coahuila.

oa a ea iter Villarreal, sp. nov. (Fig. 1). Type: MEXICO. Coanutta: Mpio.

ade Jimulco, a es verses cima 103° 13' W, 25 08°N.

eo al con Bonetiella anomal. i, Flourensia, Hechtia

Spiraea y Avalia, 1800-1850 m, 10 Ago 1994 Ja ah 7781 y M.A. Carranza.
(HOLOTYPE: MEXU; isotypes: ANSM, ENCB, TEX).

Ruellia occidentalis (Gray) Tharp & Barkley similis sed differt flores plus brevis, corolla
tubis plus brevis, calyx lobis longius quam corolla tubis; flores cleistogamous absens et
differt distributio.

Perennial herb from clustered fibrous roots; stems erect to ascending 30—
50 cm tall, densely glandular pubescent with straight hairs about 1 mm
long; the internodes 4-10 cm long; leaves with petioles 6-30 mm long,
blades broadly ovate to obovate, 3-10 cm long, 2—8 cm broad, the base
obtuse to rounded, briefly decurrent along the petiole, apex obtuse to acute,
the margin entire to undulate-crisped, both surfaces viscid with abundant
glandular hairs, the dried leaves usually green-yellowish; flowers in dicha-
sia and terminal thyrsoid panicles 5-20 cm long, to 10 cm broad, strongly

SpA 18(1): 223-226. 1998

224 Sipa 18(1)

[ tom

Fic. 1. Ruellia jimulcensis, a complete plant.

VILLAREAL Q., A new species of Ruellia 225

glandular-pubescent; peduncles ascending; calyx lobes 15—25 mm long, linear
attenuate, united at the very base, 1.0—1.2 mm broad at the base, unequal,
strongly glandular pubescent; corolla funnelform, bluish-purple 30-40 mm
long, the basal tube 8-12 mm long, the broadly campanulate throat 15—
20 mm long, the lobes 8-14 mm long, almost as broad, erose; cleistoga-
mous flowers absent; stamens didymanous, borne at the distal portion of
the corolla tube, free filaments 4-10 mm long; anthers 3-4 mm long; style
15-20 mm long; fruit ellipsoid 12-20 mm long, 3-4 mm broad, glandu-
lar pubescent; seeds 10-12, circular to oblate, 2.0—-3.0 mm long, narrowly
winged to the apex, brownish, covered with dense apressed trichomes.

Additional specimens examined: MEXICO. Coahuila: Mpio. de Torreén, Sierra de Jimulc
proximidades al ejido Trinidad, 25° 08' N, 103° 22' W, Matorral de Agave lechuguilla, Sane.
ramosa, ca er nee Jatropha y Yucca, ladera rocosa, 1900-2000 m, 25 Ago 1983,
J rreal 4387 . Carranza y A. Rodriguez (ANSM); Sierra de Jimulco, mina San
José, ne 08 N, 103° i W. Matorral desértico, 1800—1850 m, 11 Oct 1993, M.A. Carranza
1951 y J. Noriega (ANSM); ca. 54 air km SSE of Torreon in canyon above Estacion Otto in
SW side of Sierra de Jimulco near Mina San Jose, in limestone area with Acacia, Celtis,
Viguiera, geues ium, Jatropha, ee pep alli Yucca, 25° 04' N, 103° 13' W, 1850 m, 12
Sep 1980, J. Henrickson G P. Bekey 18504 (TEX). Durango: Mpio. de Cuencamé, aries El
Rosario, camin a la estacién de microondas Sapioris, carr. 49, 30 km al SE de Lerdo,

24'N, 103° 43' Ww. Matorral de Acacia crassifolia, Voie stenoloba, Bursera schlendentali, ee
es y Fouquieria splendens, 1750-1800 m, 16 Ago 1991, J.A. Villarreal 6243 y M.A.
Carranza (ANSM); Estacion de microondas Sapioris, ca. 30 km SW of Gomez Palacio on
wy to ee 25° 24°30" N, 103 43' W, matrorral desertico microfilo, 1400-1500 m,
25 Mar 1973, M.C. Johnston, T:L. Wendt & F. Chiang 10409 (TEX), just SSE of en
Microondas Sapioris, about 20 km NW of Estacion Chocolate, 25° 25'N, 103 43! W,
1500 m, 14 Aug 1973, M.C. Johnston, T.L. Wendt, F. Chiang & J. ne 12210 a
Microondas Sapioris, along cobblestone road which departs from mee 40 N of Estacion
choses ca eae air mi ae we as SW o aioe: oe 25'N, LO 2' W, Tecoma stans,
, 1300 m, 30 He 1991, M. Mayfield,
A. ae A. ae 1093 (TEX); Mexico ae 40, 25 mi SW of Lerdo, 6 nov 1964, D
Flyr 251 (TEX); Mpio. de Lerdo, 4 mi southwest of Chocolate, route 31, growing beneth
fee on clay flat in valley, 23 Jul 1958, D.S. Correll & 1M. Johnston 20008 (TEX); ca. 4
W of Ciudad Lerdo along Hwy 40 to Zacatecas, on limestone hillside with Larrea,
on Acacia, Opuntia, Agave etc., 25 31'N, 103 32' W, 1200 m, 21 Sep 1978, J. Henrickson
G E. Lee 17474 (TEX

Ruellia jimulcensis is found on sandy hillsides and dry canyons at eleva-
tions of 1200—2000 m, in xeric shrublands at the complex of mountains
near Sierra de Jimulco and Sierra del Rosario.

The new species has leaf blades broadly ovate, obtuse to rounded at the
base as R. occidentalis and often with strongly stipitate glandular trichomes
covering the stems and inflorescence. It differs in its flowers 3—4 cm long,
the basal tube of the corolla 8-12 mm long, the calyx lobes longer than the
basal tube, the cleistogamous flowers unknown and its distribution, as marked
in the key below. Tharp and Barkley (1949) gave the name R. occidentalis

226 Sipa 18(1)

var. ferrisae to a specimen from the mountains near Monterrey, Nuevo Leon
which is recognized by Henrickson and Johnston (in press) by this name
for the specimens described here. The type specimen from the Monterrey
area has oblong-obovate leaf blades, lacks stipitate glands on stem and leaves
and flower dimensions that better fit with R. nudiflora, a common species
in Monterrey area. The populations of the proposed species grow allopatri-
cally in an isolated area at the southwestern corner of the Chihuahuan Desert
Region while most of the related species are distributed in the eastern Si-
erra Madre Oriental (Turner 1991).

The three species of Rve/lia previously mentioned can be separated by
the following key:

|. Leaf blades ovate to elliptic-obovate, mostly less than 3 cm wide, sparsely to
R

moderately pubescent; lower internodes pubetulent to: ADRAC ss. cccsskaeay . nudiflora
1. Leaf blades broadly ovate to subdeltoid, 4—7(—9) cm wide, usually densely

— lower internodes on oe ee glands 2

2. Flowers (chasmogamous) 4. —6.5) cm long; basal tube of the corolla

2.5- m long, longer He = ae a cleistogamous flowers (smaller
than the chasmogamous) frecuently located in the lower nodes; se Texas,
ne Mexico (e Coah, ne N.L., Tamps, ne Ver.) R. occidentalis
. Flowers 3.5—4.0 cm long; basal tube of the corolla 8—12 mm long, shorter
than the calyx lobes; cleistogamous flowers absent; sw Coahuila and ne
Durango R. jimulcensis

No

ACKNOWLEDGMENTS
I thank Tom Wendt for the loan of specimens for revision from TEX/LL.
The illustration was prepared by Cuauhtemoc Gonzalez de Leon.

REFERENCES
HeENrIcKSON, J. and M.C. JOHNSTON. (in press). flora of the Chihuahuan Desert region.
(Ruellia). Los Angeles, California. 2:1151-11
{ARP, B.C. and F. Barktey. 1949. Genus ies in a Devas Amer. Midland Nat. 42:1—86.
TURNER, B.L. 1991. Texas species of Rvel/ia (Acanthaceae). Phytologia 71:281—299.

UNA NUEVA ESPECIE DE AGAVE, SUBGENERO
LITTAEA (AGAVACEAE) DE GUERRERO Y
OAXACA, MEXICO

ABISAI GARCIA-MENDOZA

Jardin Botdnico, U.N.A.M.
A.P. 70-614, Del Coyoacan
04510 México, D.F., MEXICO
ESTEBAN MARTINEZ SALAS
Depto. de Botdnica, Instituto de Biologia, U.N.A.M.
.P. 70-233, Del. Coyoacdn
04510 México, D.F., MEXICO

RESUMEN

Se describe e ilustra Ag “lis de los estados de Guerrero y Oaxaca, México. La especie

pertenece al grupo Striatae Baker, a ee Littaea (Tagliabue) Baker, y muestra si-
militudes con A. dasylirioides Jacobi & Bouché.

ABSTRACT
Agave gracilis from Guerrero and Oaxaca, Mexico, is described and illustrated. The spe-
cies is a member of group Striatae ES subgenus Littaea (Tagliabue) Baker. It is similar
to A. dasylirioides Jacobi & Bouc

Las exploraciones botanicas recientes realizadas en los estados de Guerrero
y Oaxaca, México, revelaron la existencia de una nueva especie de A gave, pertene-
ciente al subgénero Littaea (Tagliabue) Baker, grupo Striatae Baker, de la
familia Agavaceae. El grupo Striatae es endémico de México y los cinco taxa
que lo conforman se caracterizan por presentar hojas estriadas, lineares, con
el margen serrulado, flores campanuladas 0 cilindricas con el tubo bien desarrollado
y ovario sin cuello que se proyecta hacia el interior del tubo (Gentry 1982).

Agave gracilis Garcfa-Mend. & E. Martinez, sp. nov. (Fig. 1). Treo: MEXICO.
GUERRERO: Municipio de ras El Salado, 8 km al N de Tlapa, camino a Huamuxtitlan,
selva baja caducifolia, 990 m, 16 Nov 1982 (fls), E. Martinez et al. 2639 (HOLOTIPO:
MEXU; tsotipos: BRIT, ce K, MO).

Plantae perennes, caespitosae. Folia 40-60 x 0.4—0.9 cm, linearia, striata, margine ee,
denticulata, glauca vel viridi-glauca. Flores campanulati 2—2.2(—2.5) cm longi, tubo
mm longo, 5-7 mm lato; filamenta 2.7—3 cm, longa, in apice tubi inserta; ovar1um ais
carens, in tubi interiorem projectum; capsulae globosae 9-10 x 8—9 m

—

Plantas perennes, cespitosas, con troncos rastreros de hasta | m de largo;
rosetas hemisf€ricas, compactas, 50-80 cm de didmetro, 50—60 cm de alto.

Sipa 18(1): 227-230. 1998

228 Sipa 18(1)

LEZ
WA

SSS

— == ; yy Y} ; Fg

FAI

5 fi

IG. 1. Agave gracilis. a) planta completa con inflorescencia, b) bracteas superiores del peduinculo,
c) hoja, d) detalle del margen de la hoja, e) flores pareadas en la inflorescencia, f) flor disectada,
g) capsulas, h) semillas. Hustracién basada en los especimenes E. Martinez et al., 2639 y A.
Garcia-Mendoza y L. dela Rosa 6508.

GarciaA-MENDOZA AND MartTiNEz, Una nueva especie de Agave de México 229

Hojas mds de 100 por roseta, 40-70 cm de largo, 0.4—0.9 cm de ancho en
la parte media, ensanchandose en la base hasta 1—1.5 cm, lineares, estriadas,
planas, flexibles, glaucas o verde-glaucas, subcoridceas; margen finamente
denticulado, amarillento; espina 3—6(—8) mm de largo, débil, de color pardo-
rojizo. Inflorescencia de 1.8—2 m de largo, erecta o ligeramente inclinada,
espiga en el cuarto superior o mitad superior del pedtnculo, bracteas del
pedtinculo 5—15 cm de largo, 2—4 mm de ancho en la base, disminuyendo
en tamafio hacia el dpice, lineares, pardas, débiles, sin espina o ésta apenas
marcada; brdcteas florales 2—3.5 cm de largo, 1-2 mm de ancho, lineares,
pardas, persistentes, mas largas que las flores. Flores 2—2.2(—2.5) cm de largo,
campanuladas, verdes, con el apice de los tépalos rojizo-oscuros; pedicelos
1 mm de largo, alargdndose a 2 mm durante la fructificaci6n; ovario 7-10
mm de largo, 2-4 mm de ancho, cilfndrico, sin cuello, glabro o glabrescente,
penetrando ligeramente en el tubo del perianto, éste de 3-4 mm de largo,
5—7 mm de ancho; tépalos 0.9—1.1 cm de largo, 2.5—3.5(—4.5) mm de ancho,
oblongos; filamentos 2.7—3 cm de largo, insertos en el apice del tubo; anteras
7—9 mm de largo, 1 mm de ancho, céntricas, rojizas. Capsulas 9-10 mm de
largo, 8-9 mm de ancho, globosas, pardo-oscuras, con el perianto, estambres
y estilo de la flor persistentes. Semillas 3-3.5 mm de largo, 2—2.5 mm de
ancho, engrosadas en el lado curvo, negras.

Paratipos: MEXICO. Guerrero: Municipio de Atlixtac, 1 km al O de Santa Isabel, 30
km al E de Chilapa, carr. a Tlapa, 25 Nov 1989 (fl), J.L. Contreras 2651, 2652 (FCME);
Municipio de Tlapa, rio El Salado en su unién con el rfo Tlapaneco, 8 km al N de Tlapa,
carr. a Huamuxtitlan, 7 May 1997 (fr), A. Garcia-Mendoza y L. de la Rosa 6508, 6511,
6514 (ENCB, MEXU); Municipio de Zumpango del Rio, Xocohite, Xochipala, 23 Abr
1993 (fr), A. Gomez s.n. (MEXU). Oaxaca: Distrito de Huajuapan, El Boquerén, cafion del
tio Mixteco, 5 km al N de Tonald, 13 Ago 1993, A. Garcéa-Mendoza y F. Palma 5811 (MEXU).

La especie aqui descrita se localiza en la cuenca intermedia del rio Balsas,
en los estados de Guerrero y Oaxaca. Seguramente su distribucién se extiende
hacia otros afluentes del rio y posiblemente mas alla, siendo quiz, la planta
sefialada por Ullrich (1990) para el cerro Maouye en ee beter de Tlaxiaco,
Oaxaca. Ag crece en laderas y | re suelos derivados
de rocas Blas entre los 850 y 1300 m snm. Habita oe en
las selvas bajas caducifolias y su ecotonia con los bosques de Quercus-Pinus.
Algunos de los géneros frecuentes con los que convive son: Acacia, Agave,
Bursera, Euphorbia, Hechtia, Ficus, Forchhammeria, Neobuxbaumia y Pachycereus.

Agave gracilis presenta similitudes morfolégicas con Agave dasylirioides
Jacobi & Bouché, especie de la que se diferencia por su habito cespitoso,
roseta mas reducida en tamafio, hojas mds cortas y angostas, inflorescencia
erecta o ligeramente inclinada y flores mas pequefias; las cdpsulas globosas
y de tamafio mds reducido contrastan con las capsulas oblongas y mas grandes

230 Sipa 18(1)

Tasia 1. Comparacion de algunas caracteristicas morfolégicas entre Agave gracilis y A. dasylirioides.

Caracter Agave gracilis Agave dasylirioides

Plantas cespitosas solitarias

Tronco hasta 1m, rastrero hasta 1.5 m, rastrero

Space de la roseta 50-80 cm 100-200 ¢

‘Tamafio de hojas 40-70 X 0.4-0.9 cm tose x 2-3.8 cm

Color de hojas glaucas o verde-glaucas verdes o verde-glaucas

Inflorescencia 1.8—2 m, erecta o (1.5—)2-2.6 m, arqueada
ligeramente inclinada

Longitud de flores 2—2.2(—2.5) cm (2.6—)3—3.5 cm

Tubo de la flor 3-4 X 5-7 mm (8—)10-15 x (8-)12-16 mm

Tamanio de filamentos 2.7-3 cm 3.5—

Insercion de filamentos dpice del tubo mitad del tubo

Capsulas 9-10 — 9mm, ane (10—) 15-20 * 6-9 mm, oblongas

Semillas 3-3.5 X 2-2.5 mn 3-4 X 2.5-3 mm

de A. dasylirivides; asi mismo, las semillas son de tamafio menor en A. gra-
cilis. Una comparacion de los caracteres mas sobresalientes de ambas especies
se presenta en la Tabla 1. Las medidas de A. dasy/irivides se tomaron en parte
de Gentry (1982) y se ampliaron de acuerdo con observaciones propias. Ambas
especies habitan en la cuenca del rio Balsas, pero A. dasylirioides ocupa la
parte alta, mas humeda y fria, entre los 1900 y 2500 m snm, sobre laderas
con suelos derivados de rocas volcdnicas, en bosques de Quercus-Pinus con
Alnus, Arbutus, Clethra, Cremnophila, Garrya, Hechtia y Salvia. La época de
floracion de las dos especies se presenta en los meses de octubre y noviembre.
El epiteto especifico se refiere al cardcter esbelto de la planta.

AGRADECIMIENTOS

Agradecemos la revision del manuscrito a Raquel Galvan, Susan Verhoek,
Wendy Hodgson y Fernando Chiang. La descripci6én latina fue hecha por el
ultimo autor. La ilustraci6n es aportacién de Albino Luna.

REFERENCIAS
Gentry, H.S. 1982. Agaves of Continental North America. University of Arizona Press,

ucson.
ULiricH, B. 1990. Ein neuer Standort fiir Agave dasylirioides Jacobi & Bouché in Oaxaca.
Kakteen Sukk. 41(8):164—166.

A NEW SPECIES OF MANDEVILLA
(APOCYNACEAE) FROM JALISCO, MEXICO

JUSTIN WILLIAMS

Department of Botany
University pe : exas
Austin, TX /8713, USA.

ABSTRACT

£ fo }

g treatment of the Apocynaceae of Mexico, a routine examination
of berber an specimens revealed the following new species, Mandevilla pringle: J.K. Wail-
liams, sp. nov. The new species is distinct from other members of its alliance (sect. Torosae)
in having subsessile leaves and larger peduncles and flowers. In addition, Mandevilla apocynifoli

(A. Gray) Woodson is here presented to be a synonym of M. foliosa (Miill. Arg.) egal

RESUMEN

En la Pipa e de un prone” tratamiento de las Spo VN de Mec: un examen
de rutina de lo
Jas Williams, sp. nov. La nueva especie se eee deo otros taiemibto: desu ere

ae) por tener hojas subsésiles, y pedtinculos y flores mds grandes. Ademas, Merrell
pee nifolia (A. Gray) Woodson se presenta aqui como un sinénimo de M. foliosa (Mill.
Arg.) Hemsl.

Key worps: Apocynaceae, Mandevilla, Jalisco, Mexico

Mandevilla pringlei J.K. Willliams sp. nov., (Fig. 1). Type: MEXICO. Jatisco:
Rio Blanco, near Guadalajara, 22 Jul 1902, C. G. Pringle 11357 (HoLoryPeE: US!).

Mandevilla foliosa (Mull. Arg.) Hemsl. affinis sed foliis subsessilis petiolis 2-3 mm longis

(vice 10-18 mm in M. foliosa), pedunculis longioribus (7-10 cm vice 0.3—1.2 cm) lobis
corollae longioribus latioribusque (12-15 X 10-14 mm vice 5-8 X 2-4 mm) plantis
suffrutescentibus (vice lignosibus ramificantibusque).

Suffrutescent herbs to 0.4 m tall, stems pubescent. Leaves 3—7 cm long,
opposite, subsessile, pubescent; petioles 2-3 mm long; blades 3—7 cm long,
1.8—2.5 cm wide at middle, ovate-elliptic, apex acute, base sub-cordate,
with 2-4 a at apex of petiole on upper side. Inflorescence lateral, in-
determinate, racemose, with 2—10 flowers; peduncles 7-10 cm long, pu-
bescent; bracts 4-6 mm long, narrowly triangular to lanceolate, straight;
pedicels 10-17 mm long, pubescent, occasionally twisted. Sepals 5, basally
fused, 5-7 mm long, ca 0.5 mm wide, narrowly triangular to lanceolate,
straight, pubescent. Corolla salverform, yellow; tube 1 1-18 mm long, constricted
at mouth, lower half internally glabrous, upper half internally pubescent;
lobes 12—15 mm long, 10-14 mm wide, obovate, acuminate, occasionally
equal in length to the tube but always greater than half its length, spread-

Sipa 18(1): 231-235. 1998

Sipa 18(1)

|

Ri RRR Rin RATER tn PT

02)s

24607
PD
MAL “tt

UNITED STATES NATIONAL MUSEUM

©. G. PRINGLE
PLANTA MEXICAN A:.
L886

STATE OF JALISCO

Fao G large ®
28, firely, (902,

Fic. 1. Holotype of Mandevilla pringlei J.K. Williams.

WILLIAMS, A new species of Mandevilla yao)

ing, pubescent. Stamens 3—4 mm long; filaments ca 0.5 mm long, pubes-
cent, straight; anthers ca 3 mm long, bases sagittate with blunt lobes. Pis-
tils 7-11 mm long; style 5-7 mm long, glabrous; ovary ovoid, ca 1 mm
long, glabrous; pistil head pentagonal, 2-3 mm long. Nectaries 5, as long
as to slightly shorter than ovary. Mature follicles unknown, immature fol-
licles fused at apex, cues

Additional speci amined: MEXICO. Jatisco: Mpio. Mazamitla, Fraccionamiento
Los Cazos, to the S$ of Mazamitla, 9 Jul 1995, Machuca 7351 (TEX); Cerro Viejo, S face,
above Zapotitan de Hidalgo, a village 1 mi N of Hwy MEX 15, ca 25 mi due S, or 45 road
mi from Guadalajara, base of mountain in open jae just E of trail, ale 1890 m, 27 Jun
1956, D. P. Gregory & G. Eiten 223 (P, SMU); Huejotitan, Jul 1912, Digwer s.n. (P).

Distribution. —Mandevilla pringlei is known only from five collections
made from the Pine-Oak forest near Guadalajara, Jalisco, Mexico.

The new species honors Cyrus Guernsey Pringle (1838-1911) promi-
nent collector of the Mexican flora, and whose collections led to many a
new species in the Apocynaceae, and other families.

All specimens are with flowers and buds, Machuca 7351 is also with fruit
but the fruits are extremely immature, only 5 mm in length. The closest
relative of Mandevilla pringlei, M. foliosa (Mill. Arg.) Hemsl., has an aver-
age follicle length of 10 cm.

It should be noted that according to Pringle’s diary (Davis 1936, p. 199)
on the days prior to and after his collections made on July 22, 1902 he was
collecting numbers in the 8600 series. This, however, is in contrast to the
type collection of M. pringle: which is numbered 11357. The diary, how-
ever, does specifically state that Pringle was collecting along the Rio Blanco,
the type locality of M. pringlei, on July 22, 1902. In the numerical listing
of Pringle’s collections (ibid) the number / 1357 is used twice. One listing
is for E. apocynifolia A. Gray (= M. foliosa see below) the second is for an
unidentified species of Asclepiadaceae. In the absence of a thorough expla-
nation of Pringle’s numerical system it is reasonable to assume that the collection
number of the above type specimen is simply an error in numbering.

To date the most taxonomically thorough investigation of Mandevilla remains
Woodson’s (1933) revision. Infrageneric relations are at present moderately
unclear, however, Woodson provided both subgeneric and sectional divi-
sions which remain undisputed.

Mandevilla pringlei is a member of subgenus Mandevilla (as evidenced by
the lack of glands along the midrib of the upper surface of the leaves) where
it relates to section Torosae, evidenced by its suffrutescent habit and non-
twinning stems (Woodson 1933). Mandevilla pring/ei is most closely related
to M. foliosa, sharing with it an erect habit (opposed to the trailing habits of
M. karwinskii (Mill. Arg.) Hemsl. and M. torosa Jacq.) Woodson) and hav-

No
wo
aN

Sipa 18(1)

Fic. 2. Comparision of flowers. A. Mandevilla foliosa (Mill. Arg.) Hemsl. (V. Funk 27066,
TEX). B. Mandevilla pringlei J.K. Williams (P. Gregory & G. Eiten 223, SMU). Black bar
represents 3 cm.

ing leaves with an average length greater than 5 cm (vs. 2—5 cm of M. mexicana
(Mull. Arg.) Woodson). Mandevilla pringle: differs from M. foliosa ina number
of floral and habit characters that are contrasted below:

|. Branching shrubs to 1.5 m tall; leaves petiolate, petioles 10-18 mm long;
peduncles 0.3—1.2 cm long; corolla lobes S—8 (10) mm long, 2-4 mm wide,
up to but not exceeding half the length of corolla tube (Fig. 2a); stems, leaf
blades, inflorescence, and fruit glabrous or pubescent, but midrib of leaves
always pubescent; midrib hairs linear lanceolate 0.15—0.3 mm long (Fig.
3a); throughout Mexico M. foliosa

. Suffrutescent herbs to 0.4 m tall; leaves subsessile, petioles 2-3 mm long;
peduncles 7-10 cm long; corolla lobes 12-15 mm long, 10-14 mm wide,
half or more the length of the corolla tube (Fig. 2b); stems, leaves, inflores-
cence and fruit pubescent; midrib hairs triangular, 0.1—0.15 mm long (Fig.
3b); Jalisco M. pringlei

In the most recent keys to the species of Mandevilla (Woodson 1933, 1938),
M. pringlei keys out to M. apocynifolia (A. Gray) Woodson (Basionym: Echites
apocynifolia). In fact, Woodson (1933) cites the type of M. pringlez as a specimen
of M. apocynifolia. An examination of an isotype of M. apocynifolia (Palmer
734; HOLOTYPE: GH; 1sorype: NY!), however, reveals that it is a synonym of
M. foliosa (Ghiebreght s.n.; HOLOTYPE: G n.v., photo-holotype F! MO!). Al-
though the type of M. apocynifolia is without flowers, it is evident that the
sheet represents a specimen of M. fol/zosa due to its branching, petiolate leaves,
glabrous stems and fruits, and linear-lanceolate midrib hairs. Mandevilla
foliosa has both glabrous and pubescent-stemmed populations scattered
throughout its range. However, in the region of Jalisco where the type of
M. apocynifolia was collected the populations have consistently glabrous stems
and fruits.

Witiiams, A new species of Mandevilla 25

hats
aD
aa

Fic. 3. Scanning electron ae of Mandevilla midribs of leaf meray: showing
the length and oe of midrib hairs. A. Mandevilla foliosa (Mill. Arg.) Hemsl. (R. King &

T. Soderstrom 4632, TEX). B. Mandvilla pringle J K. Nee (Machuca 7351, TEX). White
bar on both photos represents 0.1 mm. Mi by the author using a Phillips
515 SEM (Cell Resource Center, Wave of eee Austin).

ACKNOWLEDGMENTS

I would like to thank Paul Fryxell for providing the Latin diagnosis and
to him and Billie Turner for reviewing the original manuscript. Thanks is
also extended to the curators of F, MO, NY, P, SMU, TEX and US for al-
lowing me the opportunity to observe their specimens. I would also like to
credit the staff of the Cell Research Center (University of Texas at Austin)
for allowing me access to their scanning electron microscope.

REFERENCES

s, H.B. 1936. Life and work of Cyrus Guernsey Pringle. Univ. Vermont, Burlington.

= ae R.E., JR. ss Mandevilla: In: Studies in the Apocynaceae IV. Ann. Missouri
Bot. Gard. 20:645—7
mak: 1938. Raa In: N.L. Britton et al., eds. N. Amer. Fl. 29:645-777.

236 Sipa 18(1)
BOOKS RECEIVED

FauTIN, DAPHNE GaiL, DouGtas J. FuruyMaA, and FRANCES C. JAMES, eds.
1996. Annual Review of Ecology and Systematics. Volume 27.
(SBN 0-8243-1427-1, hbk). Annual Reviews Inc., 4139 El Camino
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. $52.00. 648 pp.

There is quite a mix of articles reviewing the fields of ecology and systematics for 1996. A

total of 20 articles are presented followed by a Subject Index, Cumulative Index of Con-

tributing ea and a Cumulative Index of Chapter Titles, Volumes 23-27. The con-
tents are as follows: Early history ae progress of women ecologists: Emphasis sabe re-
search peat nen Forest Canopies: Methods, hypotheses, and future directions. Extinction

by hybridization and introgression. eee significance of resource polymo hsm
in fish, amphibians, and birds. Management of the Spotted Owl: A case history in conse
vation ae Trouble on oiled waters: lessons from the Exxon Valdez Oil Spill. Evol
tionary significance of local genetic differentiation. Rates of molecular evolution: Phylo-
genetic issues and cea Herbivory and plant defenses in tropical forests. Mechanisms
creating Community structure across a freshwater habitat eae Natural freezing sur-
vival in animals. See aphic a genetic models in conservation biology. Gene trees,
species trees, and systematics. Incidence and consequences of inherited environmental e
fects. Recruitment and the local dynamics of open marine populations. When does mor-
phology matter? Adaptive evolution of ee ane visual pigments in vertebrates.
Microbial eee Domains and Kingdoms. The g | ge: Size, shape, boundaries

and internal structure.

=~

JAMES STUBBENDIECK, STEPHAN L. Hatcu, AND CHARLES H. BUTTERFIELD. 1997.
North American Range Plants. Fifth edition. (ISBN 0-8032-9243-
0, pbk). pene o Nebraska Press, 312 North 14th Street, Lin-
coln, NE 68588-0484. (402) 472-3581. $25.00. 501 pp. Illustrated.

“The 200 species in this book were selected because of their abundance, desirability, or
noxious properties. The list of plants was developed over a nearly 40-year period by coaches
of range plant identification teams and faculty from the colleges and universities with range
management programs. The formal list is now the Master Plant List for the International
Range Plant Identification Contest sponsored by the Society for Range Management (1839
York Street, Denver, CO 80206).” Each species is treated with the following eee
Tribe, species, common name, life span, origin, and season, followed by more detailed in-
formation on Inflorescence pean Dagan Characteristics, Growth Character-
istics, Forage Value, and Habitat. An range map are provided for each plant.

Sipa 18(1): 236. 1998

A NEW COMBINATION IN MEXICAN
MANDEVILLA (APOCYNACEAE)

JUSTIN WILLIAMS

Department of Botany
University of Texas
Austin, TX 78713, U.S.A.

ABSTRACT
In preparation for a forthcoming treatment of the Apocynaceae of Mexico, it was discov-
ered that Echites holosericea of Sessé & Mog. has priority over the species Mandevilla sertuligera
Woodson and M. syrinx Woodson, accordingly Mandevilla holosericea (Sessé & Mog.) J.K.

Williams comb. nov. is here proposed.

RESUMEN

En la preparacion de un proximo tratamiento de las A le México, se descubrid
que Echites holosericea Sessé & Mog. tiene prioridad sane Mandal sertuligera “Woodson y
M. syrinx Woodson, como consecuencia se propone Mandevilla holosericea (Sessé & Mog.)

JK. Williams comb. nov.

Mandevilla holosericea (Sessé & Mog.) J.K. Williams, comb. nov. Basionym:
Echites holosericea Sessé & Mog., Fl. Mex. (ed. 2) 45. 1894. Type: MEXICO. MICHOACAN
“Ahualulci sylvis,” Oct 1787-1795, Sessé & Mogiito 5073 (HOLOTYPE: MA!; IsoTyPE: F!).

Ane ie syrinx Woodson, Ann. Missouri Bot. Gard. 19:53. 1932. Type: MEXICO.

-o: barranca of Tequila, 8 Oct 1893, Pringle 5422 ee PE: MO n.v.).
Mandev ae sertuligera Woodson, Ann. Missouri Bot. Gard. | 1932. Type: : MEXICO.
OACAN: rocky hills, one Station, 23 Jan 1907, te i 3890 (HOLOTYPE: US!).
Representative specimens. MEXICO. Jatisco: baranca near Guadalajara, Jun 1886, Pa/mer

98 (US). MEXICO: San Lucas, district Temascaltepec, 28 Nov 1933, Hinton 5274(US);

San Lucas, district pone 26 Dec 1933, Hinton 7176 (US); Dist. Temascaltepec,

Volcan, 1410 m, 9 Aug 1932, Hinton fee (US). MICHOACAN: rock-fields, Coru Sta-

tion, 15 Oct 1904, Pringle 1 oy (TEX, US-2 specimens). MORELOS: lava beds near Cuernavaca,

500 ft, 23 Jun 1896, Pringle 6329 (US). OAXACA: 5 km ieee road to Chilapa,

district of Teposcolula, 1800 m, 7 Jun 1985, Mendoza et al. 1467 (NY).

Sessé and Mocifio (1887, 1894) described 19 species of Echites and the
types for a majority of these names have not been located. In preparation of
a forthcoming treatment of the Apocynaceae of Mexico, the author studied
the collections of “Echites ’of Sessé and Mocifio during July 1995 at the Madrid
herbarium (MA), and from material borrowed from the Field Museum (F)
during December 1996 in order to locate and observe the types of their
Echites species. These observations revealed the following novelty.

The type of the species Echites holosericea Sessé and Mog. has up until now

Stipa 18(1): 237-239. 1998

- }]
Lcpriew tubule -
paacy..

.
¥

CCP Armee

/ 9 A Ant Cer | HISTORY MUSEUM | _,
NEGATIVE NO r

Pare es 41254

Fic. 1. Holotype of Echites holosericea Sessé & Mog.

HERBARIUM HoRTt E

4, (ro Plantae No
ti sé, Mocifa, ©

REAL JARDIN BOTANICO.

u

Sipa 18(1

wS

WILLIAMS, A new combination in Mandevilla 239

not been formally identified and presented. The typification of E. holosericea
has been made difficult because there are no specimens so labeled among
the Sessé and Mogcifio collections at F or MA. Many of the posthumously-
published species names of Sessé and Mocifio are not the same ones that are
written on the specimen labels (Fuertes & Fryxell 1993). Sessé and Mocifio
(1894) described the species as having opposite subcordate leaves with rusty
red-white tomentum covering all parts of the plant and white flowers with
a hairy tube arranged in a compact raceme. Only one specimen (Fig. 1) in
the Sessé and Mogifio collections matches this description. This specimen
was labeled by them as “Echites tubulosa N”. The name E. tubulosa was never
validly published by them, however, the letter “N "after the name indi-
cates Sessé and Mocifo considered this specimen to be a new species (Fuertes
& Fryxell, 1993). Since only one specimen matches the description of Echites
holosericea and has a letter “N ”on the label it is considered here to be the
holotype of E. Aolosericea.

In 1936, while the Sessé and Mocifio collections were on loan to F (McVaugh
1980), the specimen disscussed above was identified by Woodson as Mandevilla
sertuligera Woodson. This identification was later confirmed by the present
author. With the presentation of the above type it becomes evident that
the of Sessé and Mogifio name has priority over Woodson’s. A new combi-
nation is thus required and is accordingly made above.

The closest relative of Mandevilla holosericea is believed to be M. syrinx
Woodson. The two species appear to have a close resemblance to one an-
other. Indeed, there are two sheets of the same collection (Pringle 13106,
US) that Woodson annotated differently, one sheet he annotated M. sertuligera
and the other M. syrinx. Woodson (1938) keyed M. sertuligera apart from
M. syrinx by its corymbose inflorescence (vs. racemose) and sepals 4—6 mm
long (vs. 2-3 mm). Both Pringle specimens key more favorably to M. sertuligera
(now M. Aolosericea). At present I have only observed paratypes of M. syrinx
(having been unsuccessful at locating the type at MO), I am inclined, how-
ever, to recognize the two species as undifferentiated and regard them both
as a synonym of M. Aolosericea.

REFERENCES
Fuerres, J. and P.A. Fryxe.i. 1993. Types of names of New World Malvaceae of Lagasca,
Ortega and Sessé and Mogifio. Taxon 42:345—
McVaucu, R. 1980. Botanical mere of the Sessé Fil Mocifio i, —— 1803).
II. The icones florae Mexicanae. Contr. Univ. ae Herb. 14:99-14
Sessé, M. and J.M. MociNo 1887. Flora Mexicana. Méx
1894. Flora Medeaue ed. 2. México
Woopson, R.E., Jr. 1938. Apocynaceae. In: N.L. Britton et al., eds. North American flora.
New York Botanical Garden, New York. 29:103-192.

Sipa 18(1)
BOOKS RECEIVED

FautiIn, DAPHNE GalL, DouGtas J. FUruyMA, AND FRANCES C. JAMES, eds.
1997. Annual Review of Ecology and Systematics. Volume 28.
(ISBN 0-8243-1427-1, hbk). Annual Reviews Inc., 4139 El Camino
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. $60.00. 712 pp.

Volume 28 is the usual variety of papers reviewing the fields of ecology and systematics.
The following list of 25 articles reflects the diversity of contributions in this 1997 Annual
Review. Contents: Molecular population genetics of social insects. Evolution of eusociality
in termites. Evolutionary genetics and genetic variation of haplodiploids and x-linked genes.
Dissecting elena! rena patterns: Examples from the Ordovician Radiation. A com-
parison of alte itegies for estimating gene flow from genetic markers. The evolu-
tion of | eco Insect mouthparts: er the paleobiology of in-
sect feeding maas ig a Sune Echinoderm larve ig phylogeny, eae the

information y, phylogeny worth. Theoretical
and empirical examination of density-dependent selection. Toward an integration of land-
scape and food web ecology: The dynamics of spatially subsidized food webs. Settlement of

marine organisms in Ae ee richness of parasite assemblages: Evolution and patterns.
brid origins of plant species. Evolutionary genetics of life cycles. Phylogeny estimation
and hypothesis testing using maximum likelihood. Species turnover and the regulation of
trophic structure, Extinction vulnerability and selectivity: Combining ecological and pa-
leontological views. Tree-grass interactions in savannas. Plant competition underground.
Male and female alternative reproductive behaviors in fishes: A new approach using inter-
sexual dynamics. The role of hybridization and 1 ARO RFE SS Ion in the diversification of ani-
mals. The ecology of interfaces: Riparian zones. Allometry for sexual size dimorphism: Pattern
and process in the coevolution of body size in males and females.
The usual indexes follow: Subject I

ndex, Cumulative Index of Contributing Authors,
and a Cumulative Index of Chapter Titles, Volumes 24-28.

JANICK, JULES, ed. 1997. Plant Breeding Reviews. Volume 15. ISBN 0-
471-18904-9, hbk). John Wiley & Sons, Inc., 605 Third Avenue,
New York, NY 10158-0012. $145.00. 395 pp.

This volume of Plant Breeding Reviews i is dedicated to Arnel R. Hallauer, scientist, maize
breeder, and quantitative geneticist. The contents start with a dedication to Dr. Hallauer:
1) Dedication: nei R. Hallaver, ans Maize Breeder, Quantitative geneticist. T
remaining ht ter

1e€
cover a variety of areas in plant breeding. Contes tS: 2) Molecules
involved in self- seen re in flowering plants. 3
re trait loci: Separating, pyramiding, and cloning. 5) Doubled Basle‘ breeding
in cereals. ¢ t: Agronomy, genetics, and breeding. 7) Cowpea breeding. 8) Recurrent
selection in oe ean. 9) Gene action and plant breeding. All of this followed by a Subject
Index, Cumulative ee Index and a Cumulative Contributor Index for volumes 1-15

Stipa L8(1): 240. 1998

TRACHYPOGON MAYAENSIS (POACEAE:
ANDROPOGONEAE): A NEW SPECIES
FROM BELIZE

JOSEPH K. WIPFF

Pure-Seed Testing, Inc.
PO. Box 449
Hubbard, OR 97032, U.S.A.
joseph @ turf-seed.com

STANLEY D. JONES
Herbarium (BRCH)

Botanical Research Center
P.O. Box 6717
Bryan, TX 77805-6717, U.S.A.
sdjones @startel. net

ABSTRACT

eee mayaéensis, a new species from the Maya Mountains in Belize, is described
and illustrated. A key is provided to separate it from other species in Central America.
ee iiabon mayaénsis is readily distinguished from other Central American species by its
annual life cycle, height, leaf blade length « and width, and prominent and conspicuous sheath
auricles/ligules, that can reach to 6 cm in length

RESUMEN

Se aa y see una especie nueva, eee mayaensis, de las montafias Maya de
Beliz resenta una Centro-americanas. T. mayaénsis

se oe facilmente de ortras ee Centro-americanas por su ciclo de vida anual,
altura, longitud y anchura de las laminas oe sus ligulas con auriculas prominentes y
argo.

ioe

conspicuas, y que pueden llegar hasta 6 cm de

Trachypogon, a genus of approximately six species, is found in tropical and
subtropical America and Africa (Davila 1994). Two species are found in Central
America: T. spicatus (C. von Linné) K.E.O. Kuntze and T. vestitus N. Andersson.
Trachypogon spicatus {syns. = T. montufari (K. Kunth) C. Nees von Esenbeck;
Trachypogon palmeri Nash; T. plumosus (F. von Humboldt & A. Bonpland ex

n Willdenow) C. Nees von Esenbeck; and T. secundus (J. Presl) EK Lamson-
Scribner] is found from southern Texas in the United States to Argentina,
and in Africa. Trachypogon vestitus is found from Honduras to Brazil. A col-
lection of Trachypogon was made in the Maya Mountains in Belize that could
not be assigned to any known species and is here described as new.

Stipa 18(1): 241-245. 1998

242 Sipa 18(1)

Trachypogon mayaénsis Wipft & $.D. Jones, sp. nov. (Fig. 1). Typus: BELIZE.
Ayo Disrrict: 1.7 mi SE from the entrance gate inside Mountain Pine Ridge (SSW

i Belmopan) in the Maya Mountains, disturbed mountain pine forest, associates:

te Andropogon, Panicum, Axonopus, Scleria, Diodia, and Aristida; occasional, 19 Jan

14, 8.D. Jones 10489 & R. Olive (HOLOTYPE: MICH; tsoryrrs: BRCH, MO, US).

Gramen annuum caespitosum, culmis (1.59—)1.95— ae m altis; auriculis (3—)4—5(—6)
cm longis (auriculae ligulae adnatae); folia laminis (30— )43-61 cm longis; inflorescentia
9-30 cm longa.

Plants (1.59—)1.95—2.10 m tall, robust annual, densely cespitose, erect.
Leaves cauline (measurements taken from mid-culm leaves); odes ca. 11,
appressed pubescent with trichomes to 2 mm long, rooting at lower nodes;
sheaths with ascending to appressed scattered trichomes or glabrous, when
pubescent, trichomes 1.3—5.5 mm long, becoming denser at apex of sheath;
collar glabrous; sheath auricles (3—)4—5(—6) cm long, erect and adnate to the
ligules, appressed pubescent or glabrous; /zgw/es (3—)4—5(—6) cm long, mem-
branous, firm, brown, veined, appressed pubescent attenuate, adnate to the
sheath auricles; blades (30—)43—61 cm long, 6—7.5 mm wide (measurements
taken from mid-culm leaves), flat, apically long attenuate and basally long
cuneate with the basal portion becoming involute, antrorsely scaberulous
on both surfaces, margins antrorsely scabrous. L7florescence 9-30 cm long, a
spicate raceme or a panicle with two racemose branches; central axis of spi-
cate raceme or the racemose branches (if a panicle) short pubescent, intern-
odes 3—4 mm long (in middle); one pedicellate spikelet terminating spi-
cate raceme or branch. Spikelets paired, one (lower spikelet) of each pair short
pedicellate, persistent, awnless, staminate, dorsally compressed; other spikelet
(upper spikelet) of pair, longer-pedicellate, deciduous, perfect-flowered, awned,
nearly terete. Florets without paleas. LOWER (SHORT PEDICELLED)
SPIKELETS: staminate, sometimes with a vestigial ovary present; the lower
spikelet either absent or rudimentary in the lowest 4—5 pairs of spikelets in
the inflorescence or racemose branch. Pedice/s 1-2 mm long, pubescent. Spikelets
6.5—7.6 mm long, |—1.6 mm wide, pubescent; first g/umes 6.5—7.6 mm long,
1—-1.6 mm wide, 9—1 1-veined, coriaceous, partially enclosing rest of spike-
let, narrowly elliptic, sparsely to densely short pubescent, keeled, the keels
strigose-ciliate, apex bifid, the teeth 0.2-0.3 mm long; second glumes 6.5—
7.6 mm long, ca. 1.4 mm wide, 3-veined, narrowly elliptic, the margins
overlapping, ciliate; Lower Floret: lemmas 3.2-5.5 mm long; 0-veined, hya-
line, ciliate on margins; paleas absent. Upper Floret: lenmas 4.7-4.9 mm long;
3-veined, hyaline, ciliate on upper margins; pa/eas absent. UPPER (LONG
PEDICELLED) SPIKELETS: perfect flowered, awned. Pedicels 2.5—3.1 mm
long, pubescent. SprKelets (including callus) 9.5-10.6 mm long, 1-1.4 mm
wide, pubescent; ca//us, at base of spikelet, 1.8-2.0 mm lon pubescent
with trichomes to 3.0 mm long; férst g/umes 7.5—8.5 mm long, 1—1.4 wide,

WIPFE AND JONES, A new species of Trachypogon from Belize 243

f
|
i

Fic. 1. Trachypogon mayaénsis {S.D. Jones 10489 & R. Oliver (BRCH)}. A. Habit (bar equals
5 cm). B. Section of culm showing the auricle/ligule (bar equals 1 cm).

244 Sipa 18(1)

9—11-veined, coriaceous, margins involute and partially enclosing rest of
spikelet, elliptic, densely short pubescent throughout, lateral veins near apex
strigulose, pues rounded and ciliolate; second glumes 7.7-8.9 mm long, 0.9—
1.3 mm wide, 3-veined, coriaceous, narrowly elliptic, sparsely pubescent
between veins; upper margins overlapping and ciliate, apex broadly acute
and ciliate; Lower Floret: neuter (i.e. without reproductive structures); /em-
mas 6.S—7.3 mm long, 1.1-1.2 mm wide, 2-veined, hyaline, ciliate on up-
per margins, apex truncate and ciliate; pa/eas absent. Upper Floret: perfect,
lemmas 6—6.9 mm long, 0.7-1.0 mm wide; glabrous, the lower 1/3 is 3-
veined, hyaline and easily tearing, the upper 2/3 is O-veined, subcoriaceous
to coriaceous and flattened, turning into a terete awn; awn 38-50.5 mm
long, twice geniculate, pubescent from base to second bend; pa/eas absent.
Stamens 3, anthers ca. 3.8 mm long, 1-1.1 mm wide. Chromosome number
unknown.

Etymology.—The specific epithet refers to the Maya Mountain Range in Belize.

Phenology. —November—February.

Distribution.—Known from the Maya Mountains in the Cayo District of
Belize and from the state of Oaxaca in México.

Additional speci ined: BELIZE. Cayo District: Rio Privacion, Mountain Pine
Ridge, 26 Feb 1931, H.H. Bartlett 11785 (MICH). MEXICO. Oaxaca: District of Tuxtepe;
Chiltepec and vicinity, alt. ca. 200 m, 15 Nov 1941, G. Martinez Calderén 812 (MICH).

KEY TO THE SPECIES OF TRACHYPOGON IN CENTRAL AMERICA
(Modified from Davila 1994)

1. Plants robust annuals, 1.6—2.10 m . cee ce (3-)4-6 cm long;

leaf blades (30—)43—61 cm long, 6—7.5 mm wide T. mayaénsis
. Plants perennial, less than 1.5 m tall: Dera nie 2(—5) cm long or less;

leaf blades less than 30 cm long, 1-5 mm wide 2

2. Plants glabrous or rarely with basal nies sparsely pubescent; inflores-

—

rence a spicate raceme, rarely a panicle with two racemose branches ..... T. spicatus

2. Plants with sean and blades ee pubescent; inflorescence a

panicle with 3 (rarely 2) racemose branches or rarely a spicate raceme
T. vestitus
The closest relative of Trachypogon mayaénsis is probably T. spicatus, from
which it differs by the characters given in the above key; Trachypogon palmeri
Nash (=T. spicatis) is the name given to longer liguled forms of T. spicatus
in Mexico. There is also a taxon in Brazil with long ligules, 7. macroglossus
Trinius, but this taxon is perennial with very narrow, involute leaf blades.
Seeds of T. mayaénsis were planted in the greenhouse, of the five plants to
germinate, only one survived the transplanting to a larger container. Once
the plant flowered, it began to branch at the lower aerial nodes (typical of
annuals), but eventually the entire plant died and there were no new tillers
produced from rootstock. This was also observed in the field. The original

WIPFF AND JONES, A new species of Trachypogon from Belize 245

material was collected in November 1994, but when we returned to the
same site in June 1995, there was no sign of this taxon, though the other
associated perennial grasses were still present.

ACKNOWLEDGMENTS

We thank Paul A. Fryxell (TEX) and Tony A. Reznicek (MICH) for their
assistance with the Latin diagnosis and review of the manuscript; Paul A.
Fryxell (TEX) for providing the Spanish translation of the abstract and Tony
A. Reznicek (MICH) for loaning specimens of Trachypogon; and Guy L. Nesom
(SHST) thank for his initial review of the Latin description. We would also
like to thank Kelly Allred (NMSU) Gerrit Davidse (MO), W.E. Fox, III
(formerly at TAES), Gretchen D. Jones (USDA, AWPMRU) and Paul Peterson
(US) for their review and suggestions. We would like to acknowledge the
generosity of the late Royce L. Oliver (BRIT) for making the collecting trip
to Belize possible; his generosity is greatly appreciated. We would also like
to acknowledge Ben Shaw for the illustration.

REFERENCES
DaAviia, P. 1994. ee Nees. In. G. Davidse, M. Sousa S., and A.O. Chater, eds. Flora

Mesoamericana, Vol. 6, Alismataceae a Cyperaceae. Universi Nacional Auténoma de
México, México City, Meco. Pp. 380-381.

Sipa 18(1)
BOOKS RECEIVED

Hom, Leroy, Jerry Doi, Eric Hotm, JUAN PANCHO, and JAMES HERBERGER
997. World Weeds: Natural Histories and Distribution. ISBN
0-471-04701-5, cloth). John Wiley & Sons, Inc., 605 Third Avenue,
New York, NY 10158-0012, U.S.A. $195.00. 1129 pp. Illustrated.
The authors have presented most of the known biology of each species, maps of their dis-
tribution in more than 100 countries, illustrations designed to help with identification,
and an index of common names, and a bibliography of 3300 references to lead students
and researchers to further details that are contained in the procs papers. From the cover
“The culmination of four decades of global research, World Weeds presents comprehen-
sive and up-to-date information on over 100 weeds-addressing recent changes in such ar-
eas as crop tillage methods, herbicide use, and iris runoff. This monumental work,
featuring a wealth of original data from the author ovides extensive coverage of the
h entry contains a ful eal pee plus impor-
tant details on habitat fequirements and distribution, seed production, ecology, ae
ogy, crop impact, and more.” It is generously supplied with excellent line drawings

know biology of each species. Each

Hopcson, Grorrrey M. 1995. Economics and Biology. (The International
Library of Critical Writings in Economics). ISBN 1-85898-050-X,
hbk). Edward Elgar Publishing Limited, England and Edward Elgar
Company, Old Post Road, Brookfield, VT 05036, U.S.A. $189.95.
606 pp.

The International Library of Critical Writings in Economics series is an essential reference

source for students, researchers and lecturers in economics. Economics and Biology is number

50 in e are six parts each divided into several chapters. Part I: Biological

and iMecnanigel Analogies. Part II: Economics and eae Part ILI: Classical Eco-

nomics and the Darwinian Revolution. Part ed Marshall and Economic Biology.

Pare V: Evolution, Optimization and Rationality. - VI: Biology and Modern Econom-
ics. A name index follows.

Sipa 18(1): 246. 1998

TAXONOMY OF THE SPOROBOLUS FLORIDANUS
COMPLEX (POACEAE: SPOROBOLINAE)

ALAN S. WEAKLEY

The Nature Conservancy

Chifel Hill, NC-275 15-2067, 0S &

PAUL M. PETERSON

Department of Botany
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560-0166, U.S.A.

ABSTRACT

The Sporobolus floridanus complex is defined to include fiv North American species. Keys,
descriptions, distributions, illustrations and habitat inf provided for: S. curtissiz,
S. floridanus, S. pene S, teretifolius, and a new member of the complex from North Caro-
lina, South Carolina, and eastern Georgia, mee Weakley and P. M. Peterson.
A lectotype is chosen for S. floridanus. The taxonomic and ecological relationships of these
five species, as well as the related S. junceus os 5, eee. are compared and discussed.
Sporobolus curtissii, S. floridanus, S. pinetorum, S. silveanus, and S. teretifolius are all relatively

arrow endemics of various portions of the southeastern Coastal Plain. Each of these five
species is the locally dominant or codominant grass in fire-maintained pinelands with open
canopies of Pinus palustris, P. serotina, and/or P. elliottiz var. elliotti tit. The geographic distri-
butions and many ecological requirements of the S. cur S. floridanus, 8. pinetorum, and
S. teretifolius overlap, but they can be separated on a eee gradient.

RESUMEN
El complejo Sporobolus floridanus es definido para inclui ies de Norte America.
Se proporcionan claves, descripciones, distribucién, Tee bn e Scmmacien del habirat
para: S. curtissii, S. floridanus, S. silveanus, S. Wee y un nuevo miembro del complejo

de Sse del Norte, Carolina del Sur y del este de eeeric, Sporobolus pinetorum Weakley
. Peterson. Un lectotipo es aan para He floridanus. Las relaciones taxonémicas
ie de estas cinco especies, asi como de las especies afines S. junceus y S. heterolepis,
son comparadas y discutidas. Sporobolus curtissii, S. floridanus, S. pinetorum, S. silveanus, y S.
teretifolins son todas relativamente endémicas restrin poses a varias bantes< del sureste de la

llanura costera. Cada una de las c | , es el pasto | ninat
e€ pinos ae por quemas con dosel abierto de Pinus ee Pa serotina,
7 ° dint var. e//iortii. La distribuci6n geografica y muchos de los requerimientos ecol6gicos

de S. curtissit, 8. floridanus, S. pinetorum, y S. teretifolius se solapan, pero pueden ser separados
en base a un gradiente hidrolégico.

Sporobolus R. Br. is a worldwide genus of approximately 160 species oc-
curring in the tropics, subtropics, and warm temperate regions (Clayton &

Stipa 18(1): 247-270. 1998

248 Stipa 18(1)

Renvoize 1986). In the New World the genus is represented by approxi-
mately 45 species that generally occur on disturbed habirats, i.e., roadside
to open prairies and savannas (Peterson et al. 1995, 1997; Peterson et al.,
in press). Sporobolus is characterized by having spikelets with one floret, 1-
veined lemmas, fruits with free pericarps, and ligules with a line of hairs.
These characteristics also are found in two other genera, Calamovilfa (A.
Gray) Hack. and Crypsis W.T. Aiton. These three genera seem to share a
common ancestor and are the only New World members included in the
subtribe Sporobolinae (Chloridoideae: Eragrostideae).

The species of Sporobolus occurring in the southeastern United States form
a heterogeneous assemblage that may be informally divided into groups based
on characteristics of the inflorescence, spikelet, plant longevity, and gen-
eral aspect. One of these groups, characterized as long-lived, perennial, clump-
forming species, with open panicle inflorescences and relatively large spikelets,
consists of S. floridanus Chapman, S. curtissii (Vasey ex Beal) Small ex Scribner,
S. teretifolius Harper, S. silveanus Swallen, S. pinetorum Weakley & P.M. Peterson
(a new species described here), and, more peripherally, 8. janceus (P. Beauvois)
Kunth and S. Aeterolepis (A. Gray) A. Gray. In the course of conducting con-
servation, taxonomic, ecological, and herbarium studies in the southeast-
ern United States, it has become apparent that this group is poorly under-
stood and has been much confused by earlier authors, current collectors,

and field workers. As a consequence, an overlooked species has remained
unnamed. We will try to clarify this situation with a detailed discussion of
the five members of what we call the S. floridanus complex (S. floridanus, S.
curtissi1, S. teretifolius, S. silveanus, and S. pinetorum), with less detailed dis-
cussion of S. jancens and S. heterolepis.

TAXONOMIC HISTORY

Because of the general similarities of the taxa in the Sporobolus floridanus
complex, there has been widespread confusion about their circumscription
and distribution, and much erroneous information can be found in earlier
systematic treatments and in the ecological literature, and numerous
herbarium specimens are misidentified. A review of previous treatments of
the complex and its closest relatives will help define problems resolved in
this paper.

The two morphologically peripheral taxa were the first to be named.
The species currently known as S. jwnceus was described by Beauvois in 1812
in Heleochloa, and transferred to Sporobolus by Kunth in 1829. Sporobolus heterolepis
was first described as Vilfa heterolepis by A. Gray in 1835, and transferred
by Gray to Sporobolus in 1848.

WEAKLEY AND PETERSON, Sporobolus floridanus complex 249

The first of the core members of the S. floridanus complex was named by
Chapman (1860) as S. floridanus. It is fairly well characterized in his de-
scription:

“S. floridanus, n. sp. Panicle diffuse, large; spikelets (purplish) on long hair-like stalks;
glumes acute, the lower one barely shorter than the obtuse paleae, the upper a third longer;
leaves rather rigid, flat, pungent, very rough on the edges—Low pine barrens, Middle and
West Florida. September.-Culm 2-4 feet high. Leaves 1—2 feet long. Panicle 1-1.5 feet
long” (Chapman 1860, 1883, 1897).

The description, the location in Florida, and specimens collected by Chapman
at the time make the identity of S. floridanus clear. The only species occur-
ring in Florida likely to be confused with S. floridanus would be S. curtissii,
but it is contradicated by various parts of Chapman’s description, especially
the “panicle ... large,” the markedly unequal glumes, the rough-edged leaves,
and the length of the blades and the panicle. No type specimen was cited,
however, and we lectotypify S. floridanus below.

The next to be named was S. curtissii, as S. floridanus var. curtissii (Beal
1896). A year later, Lamson-Scribner elevated the taxon to specific rank (Lamson-
Scribner 1897). The distribution of S. floridanus was given as “moist pine
barrens near the coast, North Carolina to western Florida,” apparently on
the basis of G. McCarthy’s specimen at US (cited below) from Wilmington,
North Carolina, which is actually S. pznetorum. We know of no specimens of
S. floridanus from locations north of southern South Carolina.

Harper (1901) made a characteristically idiosyncratic addition to infor-
mation on the group with his collection of S. floridanus in Sumter Co., Georgia,
about which he commented “not definitely known outside of Florida be-
fore.” He elaborated on the toughness of the leaves mentioned by Chapman
and stated “I made some tests of their strength. A leaf from the first collec-
tion (no. 547) 5.5 mm. wide, not twisted, and perfectly dry, sustained a
weight of 27 pounds without breaking.”

In 1906, Harper described S. teretifolius from collections in Georgia, com-
menting that it is “a frequent and characteristic inhabitant of moist pine-
barrens in the Alramaha Grit region,” and that “it is unmistakable when
seen in the field” (Harper 1906). He provides excellent and clear character
differences between S. reretifolius, S. curtissiz, and S. floridanus, as well as an
excellent illustration of the highly distinctive leaf cross-section.

The first two editions of J.K. Small’s flora (1903, 1913) treated S. curtissiz
and S. floridanus, and gave generally accurate information about morphol-
ogy, habitat, and distribution. Sporobolus floridanus was stated as occurring
in “Georgia and northern Florida,” and S. curtissii was found in “Georgia,
Florida, and Alabama.” Small then added S. teretifolius (Small 1933). The

habitat of all three species was given as “moist pinelands, Coastal Plain,”

250 Sipa 18(1)

with distributions of “Fla. and Ga” for S. floridanus, “Fla. to N.C.” for S.
curtissii, and “Ga.” for S. teretifolins. The extension of the distribution of S.
curtiss1’ to North Carolina was apparently based on North Carolina speci-
mens of S. pinetorum misidentified as S. curtissii. In his key, Small (1933)

OQ:

istinguished the three species as follows:

Leaf-blades narrowly involute S. teretifolins
Leaf blades flat.
Pedicels appressed to the branches: leaf-blades about 1 mm wide .................

Pedicels spreading: leaf-blades 2-4 mm wide S. ee

Although this key generally serves to distinguish the three species, it has
been, in part, responsible for continued taxonomic confusion about the three
species and a fourth, S. pznetorum. The blades of S. teretifolius are anatomi-
cally oval, not involute; the blades of S. curtissi1, S. floridanus, and S. pinetorum
are flat, becoming involute when dry, either during drought conditions in
the field, or as a result of drying for herbarium specimens. The narrow blades
of S. curtissii and S. pinetornm usually fold when dried ina plant press. Specimens
of S. curtissii and S. pinetorum often have involute blades that superficially
resemble those of S. teretifolius. A second problem with the key involves the
width of the blade; both S. curtissii and S. pinetorum have blades regularly
reaching and sometimes exceeding 2 mm in width (when flat), and S. floridanus

characteristically has a leaf blade 3-10 mm wide (though the very narrow-
est blade on a plant may be as narrow as 2 mm wide). Therefore, specimens
of S. curtissti and the heretofore unnamed S. pinetorum have often been in-
correctly identified as S. teretifolins (by taking the first lead) or 8S. floridanus
(by correctly taking the second lead in the first couplet, but then incor-
rectly choosing the second lead in the second couplet by placing too much
emphasis on a blade width of ca. 2 mm).

Hitchcock (1935) treated S. floridanus (“low pine barrens, Georgia and
Florida”), 8. curtiss7i (“dry pine barrens, North Carolina, Georgia, and Florida”),
and S. teretifolius (“moist pine barrens, Georgia”). Once again, the inclusion
of North Carolina in the distribution of S. curtiss77 was apparently based on
miss identification of specimens of S. pznetorum.

Blomquist’s (1948) book on the grasses of North Carolina treated all material
of this complex in North Carolina as S. cvrtissiz, but states “According to
Swallen (1941) the North Carolina plants assigned to this species may be-
long to S. floridanus Chapm.” As will be seen below, neither species has been
documented for North Carolina; instead, North Carolina is within the dis-
tribution of $. pietorum (undescribed at the time and in some ways gener-
ally intermediate in characteristics) and S. ¢eretifolius (not collected 1n North
Carolina until 1991).

Swallen (1941) added S. s#/veanus to the group, based on material from
eastern Texas. Its distribution in Texas and the western Gulf Coastal Plain

WEAKLEY AND PETERSON, Sporobolus floridanus complex ah

of Louisiana makes it allopatric relative to other members of the S. /loridanus
complex. Swallen contrasted it with S. floridanus and S. teretifolius, yet vari-
ous characters (such as the appressed and larger spikelets) suggest a closer
relationship to S. curtissii. It also seems to show a clearer connection to S.
heterolepis than do other members of the S. floridanus complex, a relation-
ship which seems especially plausible given its proximity to S. heterolepis
and its habitat preferences for barrens with prairie affinities.

Hitchcock and Chase (1950) provided the only treatment to date with
as many as four of the species in the complex: S. floridanus (“low pine bar-
rens, North Carolina to Florida”), S. curtissii (“dry pine barrens, North Carolina
to Florida”), S. teretifolins (“moist pine barrens, North Carolina and Geor-
gia”), and S. s#lveanus (“open woods, western Louisiana and eastern Texas”).
The attribution of S. floridanus, S. curtissii, and S. teretifolins to North Caro-
lina are all based on misidentifications of S. pinetorum. This is because of the
generally intermediate morphology of S. pinetorum, and because the key was
not constructed very carefully for even the then-known members of the group.
Among the problems are that S, s//veanus (which has markedly appressed
spikelets) can only be reached by following the lead for “spikelets not ap-
pressed, the branches and pedicels somewhat spreading”; S. floridanus is separated
from S. silveanus by having glumes “about equal” rather than “unequal,”
when they actually have similarly subequal first glume to second glume
ratios; S. teretifolius is separated from S. floridanus by the accurate but often
mis-interpreted “blades terete vs. blades flat or folded” character; and S.
curtissii is separated from the others by a subjective and difficult to inter-
pret couplet about pedicel length and orientation.

Radford et al. (1964), by contrast, recognized only one species (S. seretifolzus)
in the complex as occurring in North Carolina and South Carolina. Unfor-
tunately, all material seen by them was actually the undescribed S. pinetorum.
Radford et al. (1968) added S. floridanus (alleged to occur in pinelands in
Lancaster County, South Carolina) and attributed S. teretifolius to “savannahs”
in thirteen counties in southern North Carolina and northern South Carolina.
Their key distinguished S. seretifolius from S. /loridanus by “blades terete or
subterete” vs. “blades flat or folded.” All material seen by them was actually
S. pinetorum, often with narrow folded blades so as to superficially mimic S.
reretifolius. Additionally, many of the county records are based on misidentified
specimens of Calamovilfa brevipilis (Torrey) Scribner, and likewise specimens
supposedly documenting county record distributions for Calamovilfa brevipilis
are in many cases actually Sporobolus pinetorum. In truth, both Sporobolus pinetorum
and Calamovilfa brevipilis occur (or formerly occurred) in most or all counties
in southeastern North Carolina and in the adjacent counties of South Carolina.

odfrey and Wooten’s (1979) manual of southeastern United States wetland
plants treated only S. floridanus and S. curtissii, inexplicably omitting S. teretifolius

252 Sipa 18(1)

(which occupies wetter habitats than S. cvrtiss7z) and S. silveanus (which oc-
curs in wetland situations in Louisiana, part of the geographic range of their
flora). Because of this, their treatment is of limited value in understanding
this group.

Brown (1993) addressed confusion between S. si/veanus, the most west-
ern species in the complex, and S. Aetero/epis, resulting in the deletion of S.
heterolepis trom the floras of Texas and Louisiana, and the addition of S. si/veanus
to the flora of Oklahoma.

SYSTEMATIC TREATMENT
KEY TO THE SPECIES

1, Panicle branches distinctly whorled at lower nodes S. junceus
|. Panicle branches single at lower nodes (though a few branches may be ir-
regularly approximate in pairs or threes) y
. First glume scaberulous, subulate above an expanded base; spikelets gray

to nearly black; base of plant relatively fibrous; grain spherical; plants of
rocky barrens and prairies of physiographic provinces inland from the Coastal
Plain S. heterolepis
2. First glume glabrous, linear-lanceolate to lanceolate, the base not abruptly
expanded; spikelets purplish (fading tan); base of plant smooth and hard,
made up of the indurated leaf bases; grain oblong (when present, usually
abortive); plants primarily of pine savannas and seeps of the Coastal Plain
of North Carolina, South Carolina, Georgia, Florida, Alabama, Louisiana,
and Texas, but extending further inland in southern Oklahoma and
eastern Texas 3

ass

>

3. Blades terete or subterete (wiry), 0.5—1.2 mm wide, oval in cross-
section (deeper than wide), sometimes irregularly channeled for por-
tions of their lengths (but lacking any flat portion), margins smooth,
distal portions often curling and twisted; pedicels with scattered as-
cending hairs 5. S. teretifolius
3. Blades flat, 0.8—10 mm wide, flat or V-shaped in cross-section (much
wider than deep), with free margins their entire length, margins scaberulous
(glabrous to scaberulous in S$. cwrtissii), distal portions normally stiff
and straight (note that the blades of the narrower-leaved species can
appear superficially wiry); pedicels without scattered ascending hairs,
either glabrous to scaberulous or scabrous 4

4. Lower glume usually as long or longer than the upper glume with
lower/upper ratio averaging 0.90—1.15; culms 30—-80(—90) cm tall;
panicle 10—25 cm long; pedicels 0.5-4(— : mm long, appressed, usually
shorter than the spikelet; leaves less than 30 cm long, smooth on

the margins . S. curtissi

aN

. Lower glume usually shorter than the upper ous witha denen
7 60—0.90; culms (30—)45—200(—250) cm tall; panicle
1 long; pedicels 2-22 mm or a. or appressed;
leaves m nestle more than 30 cm vee scaberulous on the margins .............00- 5
5. Pedicels appressed; lemmas 4.4—6.5 mm long, purple; anthers 3.5—
5 mm long; blades 1-2.5 mm a. and bluish-green; plants known
west of the Mississippi River 4. S. silveanus

a.

WEAKLEY AND PeTERSON, Sporobolus floridanus complex 295

5. Pedicels spreading; lemmas 34.3 mm long, purplish-brown; anthers
2—3.4 mm long; blades either wider than 2.5 mm, or dark green;
plants from east of the Mississippi River 6
6. Blades le )3-10 mm wide, bluish-green; panicles (18—)30—50
, 4-15 cm wide; lower/upper glume length ratio aver-
aging * 75—0.90; plants from southern South Carolina, Geor-
gia, eastern Alabama, and northwestern Florida ............. 2. S. floridanus
6. Blades 1.2—2(—3) mm wide, dark green; panicles 15—30 cm long,

m wide; lower/upper glume length ratio averaging 0.60—
0.80; plants from eastern North Carolina, northern South Carolina,
and eastern Georgia 3. S. pinetorum

ie ap otan ole curtissii (Vasey ex Beal) Small ex Scribn., U.S.D.A. Div.
Ag rostol. Bull. Le 142. 1897. (Figs. 1 a = Spor fl idanits Var. cur rissit
ee a Grass. N. Amer. 2:290. 1396. T U.S.A. Froripa: 1883, 1S
SM. eee ee ISOTYPE: US-556876!).

Caespitose perennials. Culms 30—80(—90) cm tall, erect, nodes all basal,
base diameter 1-2 mm, flattened; internodes glabrous. Sheaths glabrous to
appressed hairy, hairs up to 4 mm long; base shiny and endurated; margins
hyaline; summit with a tuft of hairs, hairs up to 4 mm long. Ligules 0.2—
0.6 mm long, a line of hairs. Blades 5—22(—28) cm long, 0.8—2(—2.2) mm
wide, flat to folded or involute, green, remaining green well into winter,
mostly glabrous above and below; base densely pilose on upper surface, white
to grayish hairs up to 4 mm long; margins glabrous to scaberulous. Panicles
10-25 cm long, 2—10(—13) cm wide, mostly open, contracted when imma-
ture, pyramidal to ovate; main axis glabrous to scaberulous; pulvini in axils
of primary branches glabrous or occasionally hairy; primary branches 2—
9(-10) cm long, ascending to spreading 10—80° from culm axis, not florif-
erous on lower 1/3; secondary branches mostly appressed; pedicels 0.5—4(—
8) mm long, usually shorter than spikelet, appressed, glabrous. Spikelets

3.5—6(—6.6) mm long, purplish-brown. Glumes (2.9—)3.5—6(—6.6) mm long,
fear tances ace. membranous, |-veined, equal to subequal; ratio of lower/
upper glume length 0.90—1.15(-1.33); lower (2.9—)3.5—6.2 mm long, apex

acuminate; upper 3.2—6.6 mm long, apex acuminate. Lemmas 3.4—4.5 mm

long, ovate to lanceolate, membranous, |-veined, glabrous; apex acute. Paleas

3.4-4.5 mm long, ovate, membranous, glabrous; apex acute. Stamens 3,

anthers 1.5—2.8 mm long, yellow to purplish. Grains 1.1-1.4 mm long,

fusiform, reddish-brown.

Common name.—Curtiss’ dropseed.

Distribution and habitat.—Fastern South Carolina south to central pen-
insular Florida, west to Florida Panhandle. Mesic to dry-mesic pine wood-
lands, in soils seasonally saturated at the surface or rather well-drained throughout

the year, usually under Pinus palustris, and sometimes also with Quercus spp.;
0-100 m. Sporobolus curtissti characteristically occurs in the following Na-

No
Al
aN

Sipa 18(1)

Fic. L. A-E. Sporobolus pinetorum (based on Weakley s.n., 31 Aug 1993). A. Habic. B. Ligule.
C. Spikelet. D. Pistil and stamens. E. Blade, lateral view. F. Sporobolus teretifolins (Weakley &
Schafale s.n., 16 Jul 1991). FE. Blade, transverse section. G. Sporobolus silveanus (Waller &
Bauml 3128). G. Four spikelets on a branch of the inflorescence. H, I. Sporobolus floridanus
(Curtiss 4054). H. Spikelec. I. Blade, abaxial view. J. Sporobolus curtissii (Duncan 7855). J.
Spikelet.

WEAKLEY AND PETERSON, Sporobolus floridanus complex Zo)

Fic. 2. Geographic distribution of Sporobol. tissit (A), Sporobolus pinetorum (B), and Sporobolus
teretifolius (C).

tional Vegetation Classification plant associations: Pinus palustris/Serenoa repens—
Vaccinium myrsinites| Aristida beyrichiana—S porobolus curtissii Woodland, Pinus
palustris/Quercus incana—Quercus stellata! Aristida beyrichiana—Sporobolus unceus—
Nolina georgiana Woodland; Pinus palustris—(Pinus elliottii var. elliottii)/Sporobolus
pinetorum—Aster veticulatus(Sporobolus curtissii) Woodland (Weakley et al. 1998).
Flowering July to November (or less typically at other seasons if stimu-
lated by fire).

Comments. —When fertile, 8. cwrtissi is readily distinguished by the short-
pediceled, appressed spikelets, and by the large lower glume. Vegetatively,
S. curtissii is distinctive in its short leaf blades and tendency to have gla-
brous leaf margins. Post-fire foliage tends to be narrow, stiff, and erect, while
unburned plants produce leaves that are shorter, wider, and laxer. It occu-
pies drier habitats than S. floridanus, S. teretifolius, and S. pinetorum, though
it can be found in mixed populations with all three species.

256 Sipa 18(1)

Representative specimens. FLORIDA. Alachua Co.: Gainesville, Combs 5.7. (GH). Baker
Co.: Olustee Experimental Forest, Olustee, 9 Jun 1936, Watkins 1 (US). Bradford Co.:
(FLAS). Clay Co.: 7 mi NW of Middleburg, abundant on peaty pond pine savanna, 28 Jul

1967, McDaniel 9541 (BRIT/VDB, GA). Columbia Co.: Lake City, 11-19 Jul 1895, Nash
2213 (GA, GH, NCU, US). Duval Co.: moist pine barrens near Jacksonville, 6 Aug & 13
Nov 1894, Curtiss 5181 (GA, GH, US). Manatee Co.: 1996, sane 1534 (NCU). oo
Co.: (reported in Clewell 1985). Orange Co.: Grasmere, 23 Sep 1898, Combs & Baker 112
(US). Polk Co.: 1996, McMillan 1534, (NCU). Putnam es 7 of Palatka, open field, 26
Oct 1940, Silvews 6737 (US). St. Lucie Co.: dry pine barrens, Fort Pierce, 25 June 1939,
Pie ni aueteane Taylor Co.: wet pine flatwoods, 1979, Godfrey 3 77245 (GA). GEORGIA.
Berrien Co.: Alapa Experimental Range, 2 mi S of Alapa, dominant (reported by Lemon

194 i Brantley Co.: moist open pinelands 1.5 mi W of Nahunta, 1953, Duncan 17036
(GH). Bryan Co.: Fort Stewart, 1996, McMillan 1061, 1064, 1065 (NCU). Charlton Co.:
Camp Cornelia, 1902, Ricker 932 (GH, US). Clinch Co.: pine barrens, Okifionokee [sic]
pocket, 8 Oct 1938, Ey/es 396 (DUKE). Coffee Co.: 5 mi south of Ocmulgee River, 18
May 1954, Cooley 2780 (US). Cook Co.: 2.7 mi NE of Barney on Ga. 76, 7 Nov 1993,
Sorrie 7756 (NCU). Dougherty Co.: dry sand, fossil dunes, East Albany, 30 Aug 1947,
Thorne 6441 (GA). Echols Co.: longleaf pine—palmetto flatwoods near Ga. 94, 17 Aug
1967, Clewell 2624 (FSU). Glynn Co.: 1996, McMillan 1082, 1082a (NCU). Liberty Co.:
Fort Stewart, 1996, McMi/lan 1068, 1074, 1075 5 (NCU). Long Co.: 1996, McMillan 1048,
10484 (NCU). Lowndes Co.: § of Melrose, 4 Sep 1902, Harper 1603 (GH, US). Tattnall

1996, McMillan 1081 (NCU). Todnabe Co.: 13 Dec 1993, Sorrie & Weakley 7801 (GA).

Wayne Co.: 1 mile S of Jesup, 1947, Duncan 7855 (GA, GH, FLAS, US). SOUTH CAROLINA.
Berkeley Co.: Francis Marion National Forest, 21 Jul Sorrie 8075 (NCU); locally
dominant, growing in dense and obviously quite old tussocks in gummy eee
flatwoods W of Tiger Corner, 30 May 1995, McMillan 7 "Kiellonar 1015 (NCU CH)
24 Jul 1996, McMillan 1748 (CLEMS, NCU, USCH).

|

(2 Bsc floridanus Chapm., Fl. South. U.S. 550. 1860. (Figs. 1, h,
1; 3).T U.S.A. FLORIDA. Franklin Co.: Apalachicola, Chapman s.n. (acre PE,
eee NY, bar code, 00127474!; tsotecroryre: US-998263!),

Robust caespitose perennials. Culms (40—)100—200(—250) cm tall, erect,
nodes all basal; base diameter 1.5—G mm, flattened; internodes glabrous.
Sheaths glabrous to appressed hairy, hairs up to 5 mm long; base shiny and
endurated; margins hyaline; summit with a tuft of hairs, hairs up to 6 mm
long. Ligules 0.2—0.7 mm long, a line of hairs. Blades (10—)25—50 cm long,
(2—)3—-10 mm wide, flat to folded, pale bluish-green and yellowing in age,
mostly glabrous above and below; base often sparsely hairy, hairs up to 6
mm long; margins scaberulous. Panicles (18—)30—50 cm long, 4—15 cm wide,
mostly open, contracted when immature, pyramidal to ovate; main axis scabrous;
pulvini in axils of primary branches hairy or glabrous; primary branches 4—
15 cm long, ascending to spreading 10—90? from culm axis, not floriferous
on lower 1/3; secondary branches ascending to spreading; pedicels 2-14 mm
long, usually longer than spikelet, spreading, scaberulous. Spikelets (3.7)
4—6 mm long, purplish-brown. Glumes (2.5—)2.8-5.7 mm long, linear-
lanceolate, membranous, |-veined, subequal; ratio of lower/upper glume
length (0.60—)0.75—0.90(—0.94); lower 2.5—5.1 mm long, apex acuminate;

WEAKLEY AND PETERSON, Sporobolus floridanus complex

ee ee (| ] f

eu 7
BX _» ( ° a a
nye

| ae
Sp : Be i ml i uu

,90° ae g0°

Ses | ee
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-

Fic. 3. Geographic distribution of Sporobolus floridanus (A) and Sporobolus stlveanus (B).

upper 3.7—5.7 mm long, apex acuminate to acute. Lemmas 3—4 mm long,
ovate to lanceolate, membranous, l-veined, glabrous; apex acute. Paleas 3—
4 mm long, ovate, membranous, glabrous; apex obtuse to truncate, rarely
minutely bifid. Stamens 3, anthers 2—3.1 mm long, purplish. Grains 1.7—
2 mm long, fusiform, reddish-brown.
Common name.—Florida dropseed.
Distribution and habitat.—Southeastern South Carolina south to north-
ern peninsular Florida, west to Florida Panhandle. Wet to wet-mesic pine
woodlands, in soils semi-permanently to seasonally saturated at the surface,
and even in places where water may pond for weeks, usually under Pzvus
elliottii var. elliottii, Pinus palustris, or Taxodium ascendens Brongn., and also
in seepage bogs, treeless swales, and depressional wetlands in pineland landscapes,
0-100 m. Sporobolus floridanus characteristically occurs in the following National
Vegetation Classification plant associations: Pinus palustris—Pinus elliotti vat.
elliottii/Cteninm aromaticum—A ristida beyrichiana—Sporobolus flovidanus Woodland;
Pinus palustris—Pinus elliottii var. elliottii/Styrax americanus var. pulverulentus/
Sporobolus floridanus Woodland (Weakley et al. 1998). Flowering July to November
(or less typically at other seasons if stimulated by fire).
Comments. —This is a common and conspicuous grass within its range,

258 Sipa 18(1)
and is often a dominant to codominant component of wet pinelands. The
wide leaves with distinctly bluish cast are distinctive in this species.

Representative specimens. ALABAMA. Houston Co.: (Gunn, pers. comm). FLORIDA:
Dixie Co.: flatwoods near Hines, 26 Aug 1937, West et al. 88-174 (FLAS, US). Duval Co.:

moist oe barrens, near Jacksonville, Oce 1894, Curtiss 3378 (GA, GH, NCU). Franklin
Co.: pine-turkey oak woodland {likely a wetland inclusion in or near this habitat}, 1 mile
fon bedee over Ochlochonee Bay, 14 Jul 1954, Ford 3124 (US). Gulf Co.: St. Joe, open

pineland, 5 Oct 1940, S#/vews 6498 (US). Levy Co.: Ellzen for Ellzey], flatwoods, infre-
quent, 3 Sep 1898, Combs 818 (US). Liberty Co.: Anderson 12303 (FSU). Madison Co.:
ae Res — (DUKE, FSU, GH). Wakulla Co.: St. Marks Refuge, Sorrie 7785
(NC Walton Co.: de Funiak Springs, 1898, Combs 446 (GH). GEORGIA. Baker Co.
Jones ne Research Station, Ichauway Plantation, Jack Baker Woods, 17 Aug 1 995,
McMillan, Kjellmark, & Drew 1110 (USCH). Berrien Co.: Forest Experiment Station, 5
mi NE of Tifton, open pine woods, moist se soil, 194 ay Biswell & Lemon PL-270 (FLAS,
and reported 1 in Lemon 1949). Brooks Co.: wet seepage area in upland pine woods, 1.4 mi
of Morven, 6 Sep 1967, Faircloth G Paincloth 4829 (GA, NCU). Bulloch Co.: , 20 April
1994, Sorrte 7859 (GA, NCU). Cook Co.: 4 mi SE of Adel, open pine forest, 27 Jee 1957,
Pohl 7630 (GH). Dooly Co.: e Lae drained cypress pond, on W side of railroad,
2.7 mi SSE of Unadilla, 31 Jul , Harper, Humphrey, & Duncan 16839 (GA). Echols
Co. ae pine-palmetto Dao on nthe E side of the Alapaha River, 3.6 mi S of Mayday,
Z21¢ 967, Faircloth 4978 (GA, NCU). Lanier Co.: open pine woodland alongside US
, on mi WSW of Lakeland, aes Oct 1967, Faircloth 4950 (GA, NCU). Long Co.: dis-
turbed pocosin, 6.3 mi W of 301 on Ga. 261, 6 Oct 1962, Bozeman o Radford 1964 (NCU).
Lowndes Co.: open longleaf pine forest on level terrain 5 mi E of Valdosta, 12 Oct 1963,
Duncan 22210 (GA). Sumter Co.: moist pine barrens, 31 “Aug | 900, Harper 547 (GH).
Tattnall Co.: common in slash pine flatwoods clearing, ca. 2 mi S of G enville, 11 ae
1972, Godfrey 72107 (FSU, NCU). SOUTH CAROLINA. Jasper Co.: d
to wet Prnus elliottiil P. palustris savanna on Rains series sandy loam with ety fine ce
inclusions, 28 Jun 1995, McMillan & Kjellmark 1035 (C LEMS, NCU, US

bo
No
ra

3. Sporobolus pinetorum Weakley & P.M. Peterson, sp. nov. (Figs. 1, a—
e; 2). Type: U.S.A. NortH Carouina. Cumberland Co.: Fort Bragg Army Base, Macridge
Danger Zone, ecotonal seepages and moist swales in upland Pinus palustris savannas
Weakley s.n.. 31 Aug 1993 (HoLotyPE: US!; isorype: NCU!).
AS. floridani \aminis foliorum 1.2—2(—3) mm latis atrovirentibus, inflorescentis15—30
lactis, culmis 45—120 cm altis, ratione longitudinus glumarum inferiorum/superiorum
0. ee 0.80, recedit.

Caespitose perennials. Culms (30—)45—120(—180) cm tall, erect, nodes
all basal; base diameter 1-3 mm, flattened; internodes glabrous. Sheaths
glabrous to appressed hairy, hairs up to 4 mm long; base shiny and endurated;
margins hyaline; summit with a tuft of hairs, hairs up to 4 mm long. Ligules
0.2—0.6 mm long, a line of hairs. Blades 20-50 cm long, 1.2—2(—3) mm
wide, flat to folded or involute, dark green, remaining green well into win-
ter, mostly glabrous above and below; base often sparsely hairy, hairs up to
4 mm long; margins scaberulous. Panicles 15—30 cm long, 2—6 cm wide,
mostly open, contracted when immature, pyramidal to ovate; main axis

WEAKLEY AND PETERSON, Sporobolus floridanus complex 239

scaberulous; pulvini in axils of primary branches hairy or glabrous; primary
branches 2—8 cm long, ascending to spreading 0-50° from culm axis, not
floriferous on lower 1/3; secondary branches ascending to spreading; pedicels
2—22 mm long, usually longer than the spikelet, spreading, scaberulous.
Spikelets 3.5—6.5 mm long, purplish-brown. Glumes 2.4—6.5 mm long,
linear-lanceolate, membranous, 1—veined, subequal to unequal; ratio of lower/
upper glume length (0.58—)0.60—0.80(—0.83); lower 2.44.5 mm long, apex
acuminate; upper (3.5—)4—6(—6.5) mm long, apex acuminate to acute. Lemmas
3.4-4,3 mm long, ovate to lanceolate, membranous, 1-veined, glabrous;
apex acute. Paleas 3.4-4.4 mm long, ovate, membranous, glabrous; apex
obtuse to truncate, often bifid with teeth up to 0.4 mm long. Stamens 3,
anthers 2.5—3.4 mm long, purplish. Grains 1.8—2.2 mm long, fusiform,
brown.

Common name.—Carolina dropseed.

Distribution and habitat.—Eastern North Carolina south to northern South
Carolina; and disjunct in eastern Georgia. Wet to moist pine woodlands, in
soils seasonally to semi-permanently saturated, usually under Pinus palustris
and Pinus serotina, also sometimes associated with Taxodium ascendens and
(in Georgia) Pinus elliottii var. elliottit. Sporobolus pinetorum characteristically
occurs in the following National Vegetation Classification plant associa-
tions: Pinus palustris—(Pinus elliottii var. elliottit)/Sporobolus pinetorum—Aster
reticulatus—S porobolus curtissii Woodland, Pinus palustris—Pinus elliottii var. elliottii/
Styrax americanus var. pulverulentus/Sporobolus floridanus Woodland; Pinus palustris—
Pinus serotinalSporobolus pinetorum—A vistida stricta—Erynginm integrifolium Woodland;
Pinus palustris—Pinus serotinalSporobolus pinetorum—C tenium aromaticum—E riocaulon
decangulare var. decangulare Woodland, Pinus elliottii var. elliottii—Taxodium
ascendens/ Hypericum brachyphyllum / Soonabous pinetorum—Dichanthelium scabriusculum
Woodland (Weakley et al. 1998). July to November (or less typically at
other seasons if stimulated by fire).

Comments. —This species has been overlooked until now because its over-
all morphology he it centrally in the complex, allowing it to be vari-
ously confounded with S. floridanus, S. curtissii, and S. teretifolius. Despite
its “intermediate eete it has a unique combination of characters that
warrant taxonomic recognition. Moreover, it occurs in mixed populations
with §. floridanus (in eastern Georgia), S. curtisszi (in eastern Georgia), and
S. teretifolius (in southeastern North Carolina), and in these situations the
taxa are easily distinguishable and show no signs of intermediacy or hy-
bridization. Sporobolus pinetorum is locally abundant in seasonally saturated
pinelands within its range; because it does not typically flower except fol-
lowing fire, casual field observers have often overlooked its presence and
misidentified it as Aréstida stricta.

260 Sipa 18(1)

Specimens examined. GEORGIA. Chatham Co.: pineland, Ogeechee Canal, 4 Aug 1939,
Eyles 6447 (US). Liberty Co.: in ecotone of Pinus Le ile alee beyrichiana flatwoods
and Taxodinm ascendens drain, 1995, McMillan 1967 (NCU); codominant with Sporobolus
curtissi1 in mesic longleaf pine/saw palmetto flatwoods on Fort Stewart Military
tion sec. A-5, 4 = ) 1995, McMillan, Kjellmark, & Thompson 1171 (USCH); Long Co.: in-
frequent to rare on mesic longleaf pine savanna and pond cypress depression ecotone on
Leesfield series soil transition, Fort Stewart Military Reservation on RSPAC in sect. D-12,
4 as 1995, McMillan, Kyellmark, & Thompson 1170 (UUSCH). NORTH CAROLINA. Bladen
Co.: dry savanna 10 mi N of White Lake, 8 Oct 1944, Blomquist 13617 (DUKE); flat pine
ead: 6.4 mi ESE of Ammon on road to Garland, 19 Oct 1957, AAles 37439 (NCU).
Brunswick Co.: low burned savanna on W side of road to Fr. Caswell, near junction of
Sawdust Trail, 1938, Blomquist 10448 (GH, US); burnt longleaf pine—live oak area near
inland waterway bridge, Long Beach road, 29 Oct 1950, Boyce & Godfrey 1602 (NCSC);
Southport—Supply, savanna, 23 Aug 1930, Blomquist 97 (DUKE ay Southport—Supply,
savanna, 31 Aug 1931, Blomquist 6672 (DUKE, US); Pireway, savanna, 4 Aug 1933, Schallert
s.m. (US); shrubby area in pine barrens, Boiling Springs Lakes s eee about 2 mi N of
jet. of NC 87 & 133, off rte. 133 (N of Southport), 24 Oct 1962, Terrell & Smith 3697
(NCU); plowed fire lane through a bog, Orton, 28 Sep 1941, Godfrey10053 (NCU). Co
lumbus Co.: cut over pine savanna to E of Co. road, 2.6 mi straight NW of Nakissa ae
probably Nakina], 17 Oct 1958, Bell 15739 (NCU); Schulkens Savanna, 1.7 mi W of Old
Dock, 17 Jul 1991, Weakley, Schafale, & LeBlond s.n. (NCU, US). Cumberland Co.: Fort
Bragg Army Base, Macridge Danger Zone, ecotonal seepages and moist swales in upland
Pinus palustris savannas, 31 Aug 1993, Weakley s.n. (US, NCU). Duplin Co.: transition

ear Kenansville, 14 May 1925, x. c. (NCSC); burned savanna between Clinton and Seven
Springs, 7 Jul 1946, Blomquist 13936 (DUKE); ae sae bane 0.3 mile NW of
Pin Hook, 2 Aug 1957, Ables & Leisner 33175 (NCU). Green : savanna 1.1 mi SE of

Jason, 1958, Radford 40402 (FLAS). Harnett Co.: me 192,80 7116(NCU). Hoke
Co.: 7 Oct 1991, Sorrie 5945 (NCU); 9 Oct 1991, cae ee me 25 Nov 1991, Sorrie
6101 (NCU); 28 Sep 1992, Sorrie 7007 (NCU). Jones Co.: savanna, 1.2 mi SSW of Pleas-
ant Hill, 9 Sep 1958, Radford 39935 (NCU); savanna, 3 mi SE of Pink Hill, 9 Sep 1958,
seg 39838 (NCU); savanna, 4.3 mi WNW of Hargetts Ses 18 Jul 1958, Radford
598, (GA, GH, NCU). Lenoir Co.: pine forest, near NC 11, | mile N of Pink Hill, 24
as 1957, Radford 31655 (NCU). New Hanover Co.: nen 1885, McCarthy s.n.
(US). Onslow Co.: Folkstone, 3 Aug 1939, Si/vews 4856 (US); savanna, 9.6 mi N of Hollyridge,
27 Jul 1957, Ahbles & Leisner 32690 (NCU); Camp Lejeune, 23 Sep 1990, Sorrie, Weakley,
LeBlond 5282 (GH); , 5 Sep 1991, Churchill 91-170 (BRIT/VDB); cut-over savanna about
10 mi NW of Holly Ridge toward Maple Hill, common, 16 Aug 1967, Wilbur 9432 (DUKE);
W side of US 17, 0.6 mi S of fire tower, Jun 1991, Weakley & Peet s.n. (US). Pender Co.: 30
Aug. 1991, Sorrie & LeBlond 5889 (GH, NCU). Richmond Co.: seep in powerline west of
NC 177, 12 Sep 1993, Sorrie 7679 (NCU); Sandhills Game Land, 28 Oct 1997, Sorrie 9621
(GA, GH, NCU, US). Robeson Co.: woodland border, 2 mi N of Allenton, 21 Jun 1957,
Ahles & Haesloop 29011 (NCU). Sampson Co.: pine savanna, 1.3 mi SE of junction US
421 and 701 on US 421 (SSE of Clinton), 8 Aug 1957, Ahles & Leisner 33702 (NCU). Scotland
Co.: 8.3 mi SSE of Hoffman, 11 Oct 1959, McNeely 916 (NCU); 4 Sep 1994, Sorrie 8187
(NCU). [no Co.}: in Oriente Carolina Septentrionalis, locis paludosis [in eastern North
Carolina, ie ry places}, Aug 1885, McCarthy s.n. (NCU, PH, US). SOUTH CAROLINA
Berkeley Co.: Jun 1998, Peer & McMillan 6841 (NCU). Chesterfield Co.: cleared, burned-
over sab ae in the Sandhills 1 mile W of McBee, 7 Sep 1939, Godfrey 8052 (DUKE,
GH, NCS H, US); savanna, Montrose near US 52, 29 Sep 1956, Radford 18667 (NCU);

WEAKLEY AND PETERSON, Sporobolus floridanus complex 261
sandhill, 1.3 mi SW of Patrick near US 1, 29 Sep 1956, Radford 18761 (NCU); Sandhills
National Wildlife Refuge, 16 Apr 1992, Sorrie et al. 6246 (USCH); Sandhills National
Wildlife Refuge, 1995, Pittman s.n. (US); Hudsonia Flat, 6 Oct 1993, Sorrie 7746 (NCU).

4, a Saute silveanus Swallen, J. Wash. . Sci, 31235021941. as
11 g3 3). Type: U.S.A. TEXAS. Orange Co.: 10 mi NE of Orange on U.S. 90,
Sep 1940, Si/veus 6441 (HoLoTyPE: US-1817963!; isorypEs: US-1867557!, US- sie

Densely caespitose perennials. Culms 70-120 cm tall, erect, nodes sometimes
visible, mostly basal; base diameter 1.5—4.5 mm, rounded or flattened; internodes
glabrous. Sheaths mostly glabrous to appressed hairy, hairs up to 4 mm long;
base shiny and endurated; margins hyaline; summit with a tuft of hairs,
hairs up to 4 mm long. Ligules 0.2—-0.8 mm long, a line of hairs. Blades
15-52 cm long, 1—2.5 mm wide, flat to folded or involute, bluish-green,
glabrous above and below; margins scaberulous. Panicles 21—50 cm long,
5—12(—15) cm wide, open and few flowered; pyramidal to ovate; main axis
scabrous; pulvini in the axils of primary branches glabrous; primary branches
6-20 cm long, ascending and loosely spreading 20-50? from culm axis,
not floriferous on lower 1/4—1/2; secondary branches appressed to loosely
spreading; pedicels 3-8(—14) mm long, longer or shorter than spikelet, mostly
appressed, scabrous. Spikelets 4.5—7(—7.2) mm long, purplish. Glumes 3—
7 mm long, linear-lanceolate to lanceolate, membranous, 1-veined, subequal
to unequal, ratio of ieee glume length 0.6—-0.9; lower 3—4.6 mm
long, apex acuminate; upper 4—7.2 mm long, often appeues 3-veined with
lateral folds that resemble veins; apex acuminate. Lemmas 4.4—6.5 mm long,
lanceolate, membranous, 1-veined, glabrous; margins often hyaline; apex
acuminate to acute. Paleas 4.5—6.7 mm long, lanceolate, membranous, glabrous;
apex obtuse to truncate, minutely erose. Stamens 3, anthers 3.5—5 mm long,
purplish. Grains 1.8-2.5 mm long, obovoid, laterally compressed, light brownish.

Common name.—Silveus dropseed.

Distribution and habitat.—Western Louisiana west to eastern Texas and
north to southeastern Oklahoma. Wet to mesic pine woodlands under Pinus
palustris, also in adjoining glade and barren openings, and in blackland prairies,
5-200 m. Sporobolus silveanus characteristically occurs in the following Na-
tional Vegetation Classification plant associations: Pinus palustris/Sporobolus
silveanus—Muhlenbergia capillaris —Liatris pycnostachya vat. lasiophylla Wood-
land; Sporobolus silveanus—Carex meadii Herbaceous Vegetation; Sporobolus silveanus—
Tridens strictus Herbaceous Vegetation (Weakley et al. . Flowering
July to November (or less typically at other seasons if stimulated by fire).

Comments. —There has been some confusion between this species and S.
heterolepis. Allen (1992) and Thomas and Allen (1993) reported S. s7/veanus

rom Calcasieu Parish, Louisiana, and S. heterolepis from Allen Parish and/or
Calcasieu Parish, but all specimens are actually S. sz/veanus. Similarly, both

262 Sipa 18(1)

S. stlveanus and S. heterolepis have been historically reported from eastern Texas
(Correll and Johnston 1970; Gould 1975; Johnston 1990; Hatch et al. 1990).
Brown (1993) determined all material from Texas and Louisiana to be S.
silveanus, and reported S. si/veanus as a state record for Oklahoma, a conclu-
sion followed by Taylor and Taylor (1994) and Jones et al. (1997). Sporobolus
silveanus has a bluish color in the field, and closely resembles Mahlenbergia
expansa (Poir.) Trin., with which it often grows.

Representative specimens. LOUISIANA. Allen Parish: longleaf ue woods, W of Kinder,
20 Oct 1940, Brown et al. 5717 (LSU). Calcasieu Parish: 5 mi S$ tarks, 1949, Swallen
10511(GH). OKLAHOMA. Bryan Co.: prairie near Durant, 21 bn 3, Jessee s.n. (OKLA).
PEXAS. Angelina Co.: longleaf pine He ands of the as Graham Creek Wilderness,
8.8 mi S of Zavalla on US 69 and E on FR 314, 22 Sep 1979, Nixon & Ward 9659 (ASTC).
Brazos Co.: along ere 6,12 mi Sorc ollege Station, 8 Oct ae rl 250 (TAES);

Galveston Co.: on 14th street one block W of Ave. I (FM 517), Sep 1974, Waller &
Bauml [ 28 (SBSC, mer TEX). Hardin Co. qa forest a 6 mi of Kountze,
5 ¢ 964, Gould 11028 (BRIT/SMU, TAES, TEX, US). Harris Co.: Red Bluff Road,

1.5 mi oe of SH 146, NW of Seabrook, 18 Sep ie Waller & Bauml a (GH, TAES,
TEX, US). Jasper Co.: SE of Zavalla on US 63, 1.6 mi SE of the Plum Ridge Rua. 29
Aug 1978, Marietta & Nixon 486 (ASTC, TEX). Lamar Co.: Brace Prairie” 7 mi a
Paris at intersection of highway 82 and FR 32, Collins s.v., Nov 1971 (LL, TAES, TE
Newton Co.: N of Newcon, 11 Oct 1934, Parks ex Cory +e 32 (TAES). aii:
Co.: open woods eee 10 mt NE of Orange, 30 Sep 1940, Si/vews 6441 (US). Rains Co.:
in fine sandy clay between RR and highway, 3.5 mi NW of Point, 12 Sep 1948, Shinners
10239 (BRIT/SMU). Tyler Co.: longleaf pine tak 6.5 mi E of Chester on route 1745
then left 3 mi to xeric Oligocene outcrop, 19 Oct 1967, Correl/ 35172 (LL). Van Zandt
Co.: sands, Wills Point, 15 Oct 1903, Reverchon 3484 (US).

5. Sporobolus teretifolius R. “ Harper, oo Lorrey Bot, Club 332229,
1906. (Figs. 1, ie 2). Type: U.S.A. GEORGIA: Colquitt Co.: S of Melee: 20
Sep 1902, Harper 1642 Pee pe: NY; isorype: US-431954!).

Ss
>

lon

Caespitose perennials. Culms (20—)35—80(—100) cm tall, erect, wiry, nodes
all basal; base diameter 1-2 mm, flattened; internodes glabrous. Sheaths
glabrous to appressed hairy, hairs up to 4mm long; base shiny and endurated;
margins hyaline; summit with a tuft of contorted hairs, hairs up to 4 mm
long. Ligules 0.2—0.4 mm long, a line of hairs. Blades (10—)25—54 cm long,
0.5—1.2 mm wide, tightly involute or terete, green to yellowish-green, senescing
or turning tan in late autumn, glabrous above and below; base often sparsely
hairy, hairs up to 3 mm long. Panicles 10-26 cm long, 1-9 cm wide, mostly
open to somewhat contracted when immature, narrowly pyramidal to ovate;
main axis scabrous; pulvini in axils of primary aes often hairy; pri-

mary branches 1-8 cm long, ascending to spreading 0—40° from culm axis,
not floriferous on lower 1/3; pedicels 3-18 mm long, longer than spikelet,
usually spreading, with scattered ascending hairs. Spikelets 4-5.6 mm long,
purplish-brown. Glumes 2—5.6 mm long, linear-lanceolate, membranous,
1-veined, unequal, ratio of lower/upper glume length (0.53-)0.55—0.70(—

WEAKLEY AND PETERSON, Sporobolus floridanus complex 263

0.77); lower 2—3.8 mm long, apex acuminate; upper 4—5.6 mm long, apex
acuminate. Lemmas 3.4—4.4 mm long, ovate, membranous, |-veined, gla-
brous; apex acute. Paleas 3.3-4.4 mm long, ovate, membranous, glabrous;
apex acute, Stamens 3, anthers 1.5—2.6 mm long, purplish. Grains not seen.

Common name.—Wireleaf dropseed.

Distribution and habitat.—Southeastern North Carolina south to south-
ern Georgia, west to extreme southeastern Alabama. Wet to moist pine
woodlands, under Pznus serotina, Pinus elliottii var. elliottiz, and Pinus palustris,
in soils; 1O-150 m. Sporobolus teretifolins characteristically occurs in the fol-
lowing National Vegetation Classification plant associations: Pinus palustris—
Pinus serotinal Magnolia virginiana/Sporobolus teretifolius—Carex striata
Woodland (Weakley et al. 1998) and others not yet described. Flowering
July to November (or less typically at other seasons if stimulated by fire).

Comments.—The terete leaves (well figured by Harper 1906) are distinc-
tive. Dry blades (either on dried specimens or in nature under dry field conditions)
of the other eastern species with narrow blades (S. curtissiz, 8. pinetorum) can
superficially resemble those of S. teretzfolius, and have caused confusion. Sporobolus
curtissiz, however, has much shorter blades and generally occurs in drier habitats,
and the blades of S. pinetorum have distinctly rough margins (best felt by
running ones finger along the blade towards its base, or seen at 10° or greater
magnification).

Representative specimens. ALABAMA. Houston Co.: James Hughes pitcher plant bog,
1.5—2.0 mi NE of Cottonwood, burned annually, 4 Aug 1996, MacDonald 9835 (IBE).
GEORGIA. Berrien Co.: very broad powerline on N side of route 76, about 1.7 mi SW of
Nashville, 27 Oct 1994, Sorrie s.n. (NCU). Bulloch Co.: sloping moist pine barrens, about
7 mi NW of Statesboro, 12 Sep 1954, Harper 4309 (GA). Candler Co.: about 3.25 mi W
of Metter on Stillmore Road, ecotone on E side of a N-S swampy tributary of Sams Creek,
S side of road, 22 Apr 1994, Sorrie & Stowell s.n. (NCU). Coffee Co.: rather dry pine-bar-
rens near Douglas, upper Eocene overlaid by Lafayette and Columbia, 22 Sep 1900, Harper
677 (NY). Colquitt Co.: moist bees barrens near Moultrie, 20 Sep 1902, Harper 1642 (GA,
NY, US. Cook Co.: low seepage area (pitcher plant aes alongside GA 76, 2.7 mi NE of
Barney, 21 Sep 1965, Faircloth 5 O'Neal 2994 (GA). Dodge Co.: reported without precise
location by Harper (1906). Dooly Co.: reported without precise location by Harper (1906).
Emanuel Co.: moist mown margin of GA 57 just NW of milepost 5, S side of road, at
edge of shrubby ecotone of the W side of Flat Creek, NW of Swainsboro, 6 May 1994,
Sorrie s.n. (NCU). Screven Co.: E side of US 301 at South Fork Ogeechee Creek (milepost
7), S of Sylvania, 25 Oct 1994, Sorrie s.n. (NCU). Thomas Co.: Greenwood Plantation,
Thomasville, damp peaty soil ee a Magnolia virginiana—Nyssa biflora drainage, area
burned June 1985, 23 Oct 1985, Gholson, ake Komarek, & Baker 11522 (GA). Tift
Co.: Vicinity of Irby P.O. (Cyclonera Seauiepy 28 Jul 1890, Tracy s.n. (NY) {note that ee
Tift Co. was part of Irwin Co. at the time of oa collection}. Toombs Co.: among a
colony of pitcher plants, 3 mi N of Lyons, 28 Jul 1961, Banks s.n. (GA). Turner Co. ne
age slope flanking route 32, W side of Little Sand Creek, 25 Aug 1994, Sorrie s.n. (NCU
Wheeler Co.: moist pine barrens, 2 mi W of Alamo, Plummer sandy loam, Pullen opps

881 (GA). NORTH CAROLINA. Brunswick Co.: Camp Branch Savanna remnant, just
N of SR 1335, ca. 1.2 mi E of its junction with SR 1334, back edge of savanna, near fire

264 Sipa 18(1)
ditch at ecotone to pocosiny swamp, 16 Jul L991, Weakley & Gs s,m. (US). Columbus
Co.: Old Dock Savanna, south of SR 1928, ca. 0.9 mile W of Old Dock, then south on
logging road ca. 0. 3 mi, 15 Jul 1991, W dace & Schafale s.n. is) SOUTH CAR
reorgetown Co.: pine savanna in Bates Hill Plantation, ca. 4.0 km SE of the US T01/SC
261 intersection, 5 Sep 1989, ae 0. ware H). Horry Co.: wis powerline, situated
between Mose Swamp and Grass Bay, take SC 109 ca. 2 mi NW of Baker Crossroads, then
W and intersect with Santee Cooper Power right-of-way, 22 Sep es Pittman & Jordan
s.n. (USCH). Kershaw Co.: mesic slope, SC 28-102, 2.8 mi N of SC 12, 29 Sep 1958,
Duke 2979 (NCU)

RELATIONSHIPS WITHIN SPOROBULUS

Stapf (1898), Bor (1960), Clayton et al. (1974), and Baaijens and Veldkamp
(1991) have suggested infrageneric classifications in Sporobolus based pri-
marily or strictly on Old World species, and a more worldwide classifica-
tion is still lacking. Based on possession of an open or contracted panicle,
caespitose perennial lifeform, non-whorled branch insertion, intravaginal
new shoot initiation, and first glume shorter or as long as the second Pilger
(1956) erected group four in subgenus Sporobolus. He further divided group
four into two smaller groups & & B. Sporobolus heterolepis, 8. floridanus, S.
teretifolius, and S. lasiophyllus Pilg. are members of Pilger’s “2B” group. We
agree with Pilger’s assessment that S. Aeterolepis, S. floridanus, and S. teretifolius
appear to be closely related. However, S. /aszophy//us has a few unique char-
acteristics, such as its strictly basal leaves with sheath bases that are densely
lanate and thickened, and its plumbeous to dark-brownish spikelets.

Small (1933) divided Sporobolus of the southeastern United States into
seven groups, which are not considered validly published because their rank
was not indicated. He placed 8. junceus {as “S. gracilis (Trin.) Merr.}, S. floridanus,
S. curtissit, and S. teretifolius in “Graciles,” distinguished by perennial, bunchgrass
habit, large (more than 3 mm long), purplish to brown spikelets, and glumes
unequal, the second as long as the floret. Sporobolus heterolepis was not known
so it is unknown whether he would

5)

by Small to occur in his “flora area,’
have placed this with “Graci/es” or not.

We informally recognize the S. /loridanus complex, consisting of five species
(S. curtissit, S. floridanus, 8. pinetorum, S. silveanus, and S. teretifolius) with
very similar morphological features and generally similar ecological requirements
(primarily restricted to pine savannas on the Coastal Plain). All five species
exhibit the following characteristics: caespitose perennials; basal sheaths shiny
and endurated Spies as “fire-proofing”); panicles open at maturity,
with ascending panicle branches, pyramidal to ovate, primary branches not
floriferous on lower 1/3; spikelets purplish-brown to purplish; lower glume
linear-lanceolate to lanceolate, apex acuminate; grain fusiform to obovoid.
Additionally, all five species occupy fire-maintained pinelands of the southeastern
Coastal Plain, and produce culms only following removal of foliage by fire

WEAKLEY AND PETERSON, Sporobolus floridanus complex 265

(or rarely, by mechanical disturbance) [Weakley 1998].

The S. floridanus complex appears to have its closest affinities with two
pairs of species: 1. 8. heterolepis and S. interruptus Vasey, and 2.8. junceus and
S. purpurascens (Swartz) Hamilton. Sporoblus heterolepis and S. interruptus ap-
pear to be sibling species, S. Aeterolepis being widely distributed in central
North America, and extending as an uncommon disjunct into eastern North
America, and S. interruptus being endemic in northern Arizona. They share
several characters which distinguish them from the S. floridanus complex:
spikelets plumbeous (vs. purplish fading to tan), grains globose (vs. elon-
gate and laterally flattened), scaberulous lower glumes (vs. glabrous), and
fibrous sheath bases (vs. shiny and indurated). Sporobolus heterolepis appears
to be particularly closely related to S. si/veanus, the most western of the S.
flovidanus complex, and the one most likely to have been geographically
and ecologically in contact with S. Aeterolepis in recent times. Ecologically,
S. heterolepis is a species of glades, prairies, and barrens, usually overmafic,
ultramafic or calcareous rock outcrops in the interior, “hard rock” physi-
ographic provinces of central North America (extending as a rare disjunct
east to and occasionally beyond the Blue Ridge Mountains), as well as on
loess and glacial tills.

Sporobolus junceus and S. purpurascens also appear to be siblings, S. sunceus
being a species primarily of the southeastern United States Coastal Plain
(from southeastern Virginia west to eastern Texas), but extending inland to
adjacent provinces, while S. purpurascens is distributed in southern Texas,
Mexico, the West Indies, and into tropical America. They share several characters
which distinguish them from the S. floridanus complex: panicle branches
distinctly whorled in well-marked verticils (vs. panicle branches alternate
or sometimes irregularly paired or approximate) and spikelets smaller (3.0—
3.8 mm long vs. 3.5—7.2 mm long).

DISTRIBUTION AND ECOLOGY

Sporobolus curtissii, 8. floridanus, S. pinetorum, S. silveanus, and S. teretifolius
are all relatively narrow endemics of various portions of the southeastern
Coastal Plain (Figs. 2, 3). Each of these five species is the locally dominant
or codominant grass in fire-maintained pinelands with open canopies of Pinus
palustris, P. serotina, and/or P. elliottii var. elliottii (Weakley et al. 1998). The
geographic distributions and many ecological requirements of the four more
eastern species overlap, but they can be separated based on a hydrologic gradient.
Understanding the differing but overlapping distributions and ecological
niches of the species within the group provides an important basis for the
systematic treatment.

The five species of the complex share a set of apparent adaptations to the

266 Stipa 18(1)

fire-maintained habitats in which they occur. All have the basal sheaths thickened
and cartilaginous, tightly investing the growing tips of the rhizomes, and
protecting it from damage or destruction by fire. This is a conspicuous fea-
ture of herbarium specimens (if not removed by overzealous preparation),
appearing as a stramineous, shining, thickened (almost bulbous) base of the
plant. The bases of Calamovilfa brevipilis and Calamovilfa curtissii (Vasey) Scribn.
are very similar. Species of the S. floridanus complex re-sprout quickly fol-
lowing fire, with green leaf material protruding beyond the blackened basal
sheaths within days following fire. Like other southeastern bunchgrasses
adapted to fire-maintained pinelands, including Calamovilfa brevipilis, C.
curtissit, Aristida stricta, A. beyrichiana, and Cteninm avomaticum, all five Sporobolus
species generally produce culms only after having their leaves removed by
fire, although they will sometimes flower in response to mechanical distur-

ance. For this reason, nearly all herbarium specimens have the upper por-
tions of the basal sheaths conspic uously blackened.

General exclusion of fire from much of the pineland habitat of the five
species of the S. floridanus complex, combined with their flowering only in
response to fire, means that the species are often overlooked or misidentified,
and that they are severely under-represented in herbaria despite their local
abundance. The four narrower-bladed species, S. pietorum, S. curtissii, S. teretifolins,
and S. s//veanus, are often actively or passively misidentified as other sym-
patric pineland bunchgrasses, such as Aristida stricta, A. beyrichiana, or Muhlenbergia
expansa. Entire savannas of a hundred hectares or more dominated by 8. pinetorum
’ though in reality Aristida

>

have been assumed to be “longleaf pine/wiregrass,
stricta may be completely absent. For this reason, it is important that field
biologists become familiar with the vegetative characters which allow rec-
ognition of these taxa in sterile condition.

The complex reaches its greatest diversity in the Coastal Plain of Geor-
gia, where S. floridanus, S. curtissii, S. teretifolius, and S. pinetorum are all found,
and co-occur in various combinations of two and three species, along with
the related S. janceus. Sporobolus silveanus is the only species of the complex
that is fully allopatric, and the only one that occurs west of the Mississippi
River. In southeastern North Carolina, 8. pinetorum and S. teretifolins occur
in wet pinelands dominated by mixtures of Pinus palustris, Pinus serotina,
and Taxodium ascendens. They generally co-occur with other savanna bunch-
grasses, notably Creniam aromaticum, Calamovilfa brevipilis, Mublenbergia expansa,
and Aristida stricta. Sporobolus teretifolius is restricted to the wettest pine sa-
vannas, usually so wet as to exclude Aristida stricta, and it may be the sole
dominant, or codominant with S. pinetorum, Cteninm aromaticum, and Mublenbergia
expansa. Slightly less wet savannas have varying mixtures of S. pinetorum,
Aristida stricta, Muhblenbergia expansa, and Cteninm aromaticum. A similar

WEAKLEY AND PETERSON, Sporobolus floridanus complex 267

composition, though often with substantial Calamovilfa gees as well, is
seen in sandhill/pocosin ecotones in the inner Coasta

A general hydrologic gradient of the five species ue = sympatric S.
junceus would be (from wetter to drier) S. floridanus, S. teretifolius, S. pinetorum,
S. silveanus, 8. curtissi1, 8. gunceus. Sporobolus floridanus is restricted to habi-
tats which are saturated at least seasonally (and often semi-permanently)
and may even have shallow ponds for weeks or even months. The other spe-
cies are generally intolerant of ponds for more than a few days. Sporobolus
teretifolius occurs in habitats with semi-permanent to seasonal saturation,
whereas S. pinetorum occurs in habitats with seasonal saturation. Sporobolus
silveanus and S. curtissit occur in a range of sites, from seasonally saturated
to sites which are rarely saturated at the surface for periods of short dura-
tion only, and may range up to rather well-drained (though not xeric) sites.
Sporobolus junceus strictly occupies well-drained sites, ranging into truly xeric,
sandhill situations, where it reaches its greatest abundance.

Sprobolus floridanus generally occupies the wettest habitats of the com-
plex. Particularly towards the edges of its distribution, it is generally found
in very wet situations, often where water stands for periods of time, often
in seepage bogs or swales, and generally associated with Pznus ef/zottiz var.
elliottii and Taxodium ascendens. Near the center of its distribution, especially
in the eastern portion of the Florida panhandle, S. floridanus ranges into
less wet habitats, and occurs in “mesic flatwoods.” Where its distribution
overlaps with S. pinetorum, S. curtissii, and S. teretifolius, it can occur (with
clumps side by side), but it clearly ranges ecologically into wetter sites and
avoids drier sites.

The four eastern species in the S. floridanus complex commonly co-occur
in pineland landscapes, and sometimes occur in mixed populations, with
individuals of various species intermixed. In these circumstances, the vari-
ous species are always readily distinguishable by morphological characters.
No intermediates or likely hybrids have been seen. None of these species
has been investigated cytologically.

ACKNOWLEDGMENTS

We wish to thank Michael P. Schafale, Richard LeBlond, Bruce A. Sorrie,
Brian van Eerden, Harry LeGrand, Jr., Patrick McMillan, and Robert K.
Peet for discussions regarding this group and for providing additional dis-
tribution information; Susan C. Escher for preparing the illustration; Alice
Tangerini for preparing the distribution maps; Victoria E. Batista for pre-
paring the Spanish abstract; Dan H. Nicolson for help preparing the Latin
diagnosis; Dieter C. Wasshausen and Gene Rosenberg for help translating
German; the curators of the listed herbaria, and especially Jim Massey at

268 Sipa 18(1)

NCU for providing facilities to the first author; and David W. Hall, Stephan
L. Hatch, and Robert K. Peet for critically reviewing the manuscript.

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CARYOPSIS MORPHOLOGY OF LEPTOCHLOA
SENSU LATO (POACEAE, CHLORIDOIDEAE)

NEIL SNOW!
Department of Biology
Washington University

Campus Box 1137
Sa on, MO O5130, U.S.A:

P.O. Box 299
Missouri Botanical Garden
St. Louis, MO 63166, U.S.A.

ABSTRACT

Caryopsis morphology of the grass genus Leprochloa was studied regarding the necessity
of some authors for segregating Diplachne. The data do not support the separation of Diplachne
from Leptochloa based on a putative bimodal distribution of dorsal and lateral cross-sec-
tional compression. The presence or absence of a prominent sulcus and the relative adna-
tion of the pericarp are the only attributes sufficiently distinct to warrant use as phyloge-
netic markers. However, variations in surface texture and color can be useful regionally as
diagnostic characters at the species level.

Key worbs: caryopsis, Leptochloa, Diplachne, morphology, systematics, Poaceae, Chloridoideae.

RESUMEN

Se estudio la morfologia del cariépside del género Leptochloa dada la supuesta necesidad
de algunos autores de segregar Diplachne. Los datos no apoyan la separacion de Diplachne y
Leptochloa basada en una supuesta distribucién bimodal de la compresi6n dorsal y en secci6n
transversal lateral. i ede O ausencia a de un surco prominent y la adnacion relativa
del pericarpo son rentes para justificar su uso como
marcadores Bieeen cia. Sin a. las variaciones en textura y color de la superficie
pueden ser utiles regionalmente como caracteres diagnoésticos a nivel especifico.

INTRODUCTION

The genus Leptochloa P. Beauv. s.1. (including Diplachne P. Beauv.) has been
the subject of numerous regional systematic studies due to its wide geo-
graphic distribution and the relative abundance of herbarium specimens
(Hitchcock 1903; Parodi 1927; McNeill 1979; Lazarides 1980; Phillips 1982
Nowack 1994; Nicora 1995). These authors (and others) have disagreed as to
whether Diplachne should be segregated from Leptochloa, thereby mirroring

‘Current Address: Department of Biological Sciences, University of Northern Colorado,
Greeley, CO 80639, U.S.A. Email: nsnow@bentley.unco.edu

Stipa 18(1): 271-282. 1998

272 SIDA 18(1)

the differing opinions of two worldwide generic summaries of grasses (Clayton
& Renvoize 1986, Watson & Dallwitz 1992). A frequently cited source of
evidence to support the segregation of Diplachne has been differences in caryopsis
features.

Parodi (1927) apparently was the first to examine cross-sectional shapes
critically. He partitioned four neotropical species into Diplachne or Leptochloa
based on relative compression (dorsal or lateral) of the caryopsis, and the
presence or absence of a hilar groove. He suggested that Leptochloa ets
was aberrant in Leptochloa because of its lack of a hilar groove (Parodi, l.c.).

Valls (1978) studied the systematic affinities of Leptochloa dubia in rela-
tion to the generic boundaries of Leptoch/oa. He illustrated cross-sectional
profiles and profiles from the embryonic and hilar sides for seven species.
The figures revealed a gradation in cross-sectional profile from nearly round
in Diplachne caudata to somewhat triangular in Leptochloa virgata and L. scabra,
to relatively flattened in L. fascicularis (Valls, l.c.: 103). He also concluded
that a hilar depression was a tenuous systematic feature. Despite somewhat
limited sampling, his Its suggested strongly that caryopsis features intergrade
too thoroughly to split Leptochloa toa oicnousiys into two genera, although
he acknowledged that “some grouping of species can be achieved on the
basis of caryopsis type” (Valls, l.c.: 105). Unfortunately, his results were
never formally published and have not been cited by subsequent authors.

McNeill (1979: 401) and Nicora (1995: 233) repeated almost verbatim
the observations of Parodi (1927) Maes eoueae additional data or cit-
ing the work of Valls (1978). Lazarides (1980) observed that Australian species
generally could be segregated into aay or Diplachne on the basis of
caryopsis shape, with the exception of Leptochloa digitata, with its flattened
shape. Phillips (1982: 144) agreed with Parodi (1927) regarding the util-
ity of caryopsis shape for splitting the genera, but noted exceptions in Diplachne

candata, Leptochloa obtusiflora, and L. longa. Nowack (1994) provided a cur-
sory review of caryopsis shapes for Malesian taxa and concluded that the
differences set forth by Parodi (1927) were insufficient to permit recogni-
tion of segregate genera.
Based on a recent monographic treatment (Snow 1997), Leptochloa (in-
cluding Diplachne) represents forty taxa, with one species, L. monticola Chase,
eing of dubious inclusion (Valls 1978; Snow 1996). Prior to this study,
relatively few taxa had been examined critically for variation in features of
the caryopsis, and much systematic weight had been placed on the meagre
observations that existed. In light of lingering debates about generic boundaries
between Leptochloa and Diplachne (Jacobs 1987) and the emphasis previous
authors placed on the utility of the caryopsis to segregate these genera, a
survey of all currently recognized taxa of Leptoch/oa was undertaken to evaluate
whether features of the caryopsis could be useful as phylogenetic markers.

SNow, Caryopsis morphology of Leptochloa pa ie.
MATERIALS AND METHODS

Caryopses of all currently recognized taxa in Leptochloa (Snow 1997) were
removed directly from herbarium specimens (Appendix 1), placed under a
Nikon SMZ-U dissecting microscope with camera lucida attachment, and
the profiles were traced by hand. Cypholepis yemenicus was included because
it resembles L. e/ewsine and L. obtusiflora in several respects (Snow 1996),
and was used as an outgroup in preliminary cladistic studies of Leptochloa
(Snow 1997; see also van den Borre & Watson 1997). In most cases a mini-
mum of three specimens were examined for variation (Appendix 1). The
following features were observed: 1) caryopsis shape when viewed from the
hilar side (“hilar profile”); 2) caryopsis shape when viewed from a cross-sec-
tion taken at midpoint with the hilar side oriented above (“cross sectional
profile”); 3) the presence or absence of a sulcus or other depression on the
hilar side when viewed in cross-section; 4) ornamentation on the outer coat
(perisperm); 5) relative adnation of the perisperm to the endosperm, and 6)
color of the grain. To standardize the sampled developmental stage, cary-
opses were selected from spikelets in which florets were beginning to disar-
ticulate, a condition that assures their maturity. In virtually all cases the
caryopses from the lowermost floret in the spikelets were selected.

For the sake of precision, descriptive terminology of shapes follows that
of the Systematics Association (1962), whereas that of surface ornamenta-
tion follows Murley (1951). Given that shape is a continuously varying character,
these typologies might not account for the observed and often subtle varia-
tions in shape. For example, a caryopsis might have an intermediate ellip-
tic shape of 2.5:1, which is absent from the diagram. Nonetheless, after
initial analyses, the diagram shapes appeared adequate to standardize an
summarize the majority of both hilar profile and cross-sectional shapes. With
respect to cross-sectional shapes, I accounted for the absence of sharp edges
by prefixing the terms “obtriangular” (3:2), “shallowly obtriangular” (3:2),
and “shallowly obdeltate” with the word “rounded,” which more accurately
depicts their shapes. To account for concave inflections of the hilar surface
(always oriented above, Fig. 2) I used the terms “sulcus” and “depression,”
depending on the degree of concavity. As used here, a sulcus is a vertical or
nearly vertically walled groove; a depression refers to any gradual concav-
ity, and will be further modified by the terms shallow, moderate, deep, narrow,
and broad. These subjective terms were deemed necessary to describe the
observed variation.

RESULTS

The hilar profile for most species was some variation of obovate or ellip-
tic (Fig. 1; Table 1). Only four taxa had ovate hilar profiles (L. fusca subsp.

Sipa 18(1)

Narrowly elliptic

Obovate

254 32

Elliptic

8) &)

221 a2

Ovate

Widely obovate

6:5

Widely elliptic

6:5

Very widely obovate

1:1

Fic. 1. Hilar profiles of caryopses observed for Leptochloa. The apex is oriented above.

SNow, Caryopsis morphology of Leptochloa 275

Tas_e 1. Variation in hilar profiles and cross-sectional shapes of the caryopsis in Leptochloa s. 1. The
numbers following each taxon reflect observed variation: numbers to the left of the double bar (||)
indicate hilar see ae (see Fig. 1); fe to the right of the double bar reflect cross-sectional
shapes (Fig. 2).. Her 8 || 2,9 would indicate a taxon having narrowly elliptic (3:1), elliptic
(2:1), and obovate CS re se profiles, and oblate and depressed obovate (2:3) cross-sectional shapes.
Taxa with an asterisk (*) indicate those previously placed in Diplachne by some authors.

Leptoc hloa aquatica 3,9,10 || 8,9,12 L. marquisensis 2,3,6 \| 9,10

L. caudata 8 || 2 L. monticola* 3,8 || 4,8

L. chinensis 8, S| 2,9 L. nealleyi 4,9 | 2,9

L. chloridiformis 2 || 7 L. neesii 5,10 || 1

L. coerulescens 8,9 || 2,8 L. obrusiflora 3 || 8

L. decipiens subsp. asthenes 3,8 || 2,8,9 . panicea gubsp: brachiata 3,4 || 10,13
L. decipiens subsp. decipiens 3 || 9 i. panicea subps. mucronata 3 || 1,2,8
L.d 3,9 L. panicea subsp. panicea 7 || 1,2

L. digitata 2,3,8 || 8,12 L. panicoides* 3,4 || 8

L. divaricatissima 3 || 2,9 L. rupestris 3 || 10

L. dubia* 2,3,11 | L. scabra* 2, |

L. eleusine* 9 || 8, L. southwoodii 3,4,8 || 1,2

L. fusca uo cca 8 || 4 L. squarrosa 2 || 5,1

L. fusca subsp. fusca* 9 || 4, L. srilankensis 3 I 9,10,11

L. fusca susbsp. muelleri* y 5 || 4,8 L. tectoneticola* 8

L. fusca subsp. uninervia* 3,7,8 || 4 L. uniflora 2,6 || 6,

L. gigantea* 3,6,8 || 8 L. virgata 2,3,6 || y Ai led bezrdes)

L. longa 3,8 || 9 L. viscida* 3,8 || 4

L. ligulata 3,4 || 2,7 L. xerophila 3 || 2

L. malayana* 3 || 9,11 Cypholepis yemenicus 4 || 8,9

uninervia, L. gigantea, L. uniflora, L. virgata). The widest was the very widely
elliptic shape of a few specimens of L. dubia. The thinnest was the narrowly
elliptic (3:1) shape expressed by some specimens of L. caudata, L. chloridi-
formis, L. dubia, L. digitata, L. scabra, L. squarvosa, L. uniflora, and L. virgata
(the lattermost sensu lato, including L. barbata and L. procera sensu Nicora
1995). Many species were variable, for example having both elliptic (2:1)
and obovate (2:1) shapes. Not surprisingly, the greatest variation in hilar
profile shape occurred in widespread species such as L. dubia and L. virgata.

The cross-sectional shape was considerably more variable than hilar profile
shape (Fig. 2). Overall, the observed variation ranged from dorsally com-
pressed through circular (no compression) to laterally compressed (Fig. 2).
Most taxa had only slight to moderate degrees of lateral or dorsal compres-
sion. As with hilar profiles, many taxa showed infraspecific variation in cross-
sectional shapes (Table 1). Some specimens of L. weesi# appeared circular,
whereas others were oblate (Table 1). The greatest degree of dorsal com-
pression was expressed by the depressed obovate (1:2) and transversely el-
liptic (1:2) shapes. With some modifications, these shapes accounted for
some or all of the variation of many species (Table 1). The highest degrees
of lateral compression were the obovate (3:2), rounded shallowly obdeltate

276 Stipa 18(1)

(5:6), and rounded shallowly obtriangular shapes. Only L. sqwarrosa was obovate
(with a moderate hilar depression), whereas some representatives of L. virgata
were both rounded shallowly obdeltate (5:6) or rounded shallowly obtriangular
(233);

A distinct sulcus was present only for L. rupestris and L. uniflora, although
a number of taxa had depressions of varying extent on the hilar surface (see
Discussion).

The surface of the pericarp varied from smooth to variously rugose. The
following were at least occasionally somewhat rugose: L. chloridiformis, L.
decipiens subsp. decipiens, L. divaricatissima, L. gigantea, L. longa, L. malayana,
L. monticola, L. nealleyi, L. neesii, L. obtusiflora, L. scabra, L. southwoodi, L.
uniflora. \ hereas a smooth pericarp was consistent for many taxa, those that
expressed the rugose condition did so irregularly.

Species with a weakly adnate pericarp (detaching soon after placement
in water at room temperature) included L. chloridiformis, L. dubia, L. elensine,
L. fusca subspecies fusca, fascicularis, uninervia, and L. obtusiflora.

The color of the caryopsis varied from very light brown to dark reddish
or very dark brown, but most were an intermediate shade. Leptochloa longa,
L. obtusiflora, and L. squarrosa were usually dark brown. Leptochloa monticola,
a species of dubious inclusion in the genus (Clayton & Renvoize 1986; Valls
1978; Snow 1996, 1997), was usually a dark reddish brown.

DISCUSSION

This simple study of caryopsis morphology has revealed more variation
within and between taxa of Leptochloa than previously recognized (Parodi
1927; McNeill 1979; McVaugh 1983; Nowack 1994; Nicora 1995). With
two exceptions, features of the caryopsis appear to be of little value in Leptochloa
as phylogenetic markers, although some are of diagnostic value in keys. These
results contrast with those of a recent study in Triticeae, which suggested
caryopsis morphology was of systematic value at the tribal level (Terrell &
Peterson 1993).

Parodi’s (1927) study was limited to four species in Leptoch/oa and one in
Gouimia Fourn., which represents only ten percent of Leproch/oa as currently
circumscribed (ease 1997). Another disconcerting aspect was his lack of
voucher specimens and uncertain depth of sampling within taxa, alchough
this study does not contradict the profiles of the species he illustrated. Overall,
Parodi’s sampling underestimated considerably the variation in cross-sec-
tional shape in Leptoch/oa. This study has revealed nearly continuous varia-
tion in cross-sectional shape, from dorsally compressed and non-compressed
(circular or oblate) to various degrees of lateral compression. The bimodal
compression (lateral or dorsal) of caryopses in Leptochloa recognized by Parodi

SNoW, Caryopsis morphology of Leptochloa 277

Circular Oblate Transversely elliptic

OO Oe

Obovate Widely obovate Very Widely obovate
oe2 6:5 1:1
Depressed obovate Widely depressed obovate
12 2:3 5:6

"Rounded" "Rounded" shallowly "Rounded" shallowly

obtriangular obtriangular obdeltate

Y YP

Fic. 2. Cross sectional shapes of caryopses observed for Leptochloa. The hilar side is oriented

above.

278 Sipa 18(1)

(1927) simply does not exist for the genus as a whole. As such, variation in
caryopsis cross-sectional morphology cannot by itself be invoked as a ge-
neric-level character to segregate Diplachne.

The caryopsis profile as viewed from the hilar side ranges continuously
from narrowly elliptic through ovate to obovate and very widely obovate
(Fig. 1). Whereas the extremes of variation can be useful as diagnostic fea-
tures between some taxa, the continuous variation makes the hilar profile
useless as a phylogenetic marker (Stevens 1991).

Various degrees of concavity occur on the hilar side. Broad, shallow de-
pressions (not illustrated) characterize certain taxa fairly well, such as Leptochloa
eleusine, L. longa, and L. obtusiflora. Taxa showing this feature irregularly were
Leptochloa chinensis, L. dubia, and the related Cypholephis yemenicus. Shallow,
relatively narrow depressions (not illustrated) occur in other taxa, although
less consistently; these included L. chloridiformis, L. decipiens subsp. decipiens,
L, digitata, L. squarrosa, and L, virgata. At best, the degree of concavity is
useful only as a diagnostic character in regional keys. However, a promi-
nent sulcus, characterized by its vertical or nearly vertical walls, was a con-
sistent character for L. rupestris and L. uniflora, and is one of only two char-
acters I consider sufficiently distinct and consistent to be phylogenetically
useful.

The relative adnation of the pericarp is the second character of the cary-
opsis useful for phylogenetic inference. It is well known that the pericarp is
only weakly adnate to the endosperm in some species of Leprtochloa (Izaguirre
& Laguardia 1987; Watson & Dallwitz 1992) and some related genera, such
as Eragrostis Wolf (Lazarides 1997). In such taxa the pericarp will dissociate
from the endosperm quickly when placed in water at room temperature.

Most species in Leptochloa have a smooth outer texture. A few can be rug-
ose, although this feature was unreliable within taxa. For example, a coarse
but sparsely rugose surface generally, but not always, characterizes Leptochloa
panicea subspecies panicea and mucronata (sensu Snow 1998a, but not sensu
Nowack 1994), which helps to distinguish these from the widespread L.
panicea subsp. brachiata (Snow 1998a; formerly known as L. filiformis or L:
mucronata {Snow & Davidse 1993}).

s the color of the caryopsis often varies with the degree of maturity,
only mature specimens should be evaluated for this attribute. Except as a
diagnostic feature in keys, in which a few species are dark brown, color is of
minimal systematic value in Leptoch/oa.

I return now to cross sectional shape, which has been discussed exten-
sively regarding the separation of Diplachne from Leptochloa (Parodi 1927;
McNeill 1979; Phillips 1982; Nicora 1995). As mentioned above, the per-
ceived bimodality of lateral and dorsal compression discussed by Parodi (1927)

SNow, Caryopsis morphology of Leptochloa Zs)

has been invoked to segregate Diplachne from Leptochloa. The results of this
study firmly reject such a notion, given the nearly continuous variation of
cross-sectional hilar profiles (Fig. 1).

It also has been suggested that the presence or absence of a distinct keel
on the lemma is positively correlated with cross-sectional shape, and is a
means by which the genera can be separated (Parodi 1927; McNeill 1979;
Nicora 1995). It is true that some taxa with a dorsally compressed cary-
opsis have flat lemmas at maturity, as for example L. fusca subsp. muellert.
However, others have little or no such positive correlation. Mature fruits of
L. neesii can be round or nearly so in cross section, yet still be borne within
a keeled lemma (e.g., Langfield 285, CANB). This is also true for some specimens
of L. ligulata, L. nealleyi, L. panicea subsp. panicea, and L. southwoodi1. Pre-
liminary cladistic studies have failed to consistently group together taxa
having dorsally flattened caryopses (Snow 1997).

This study upholds and strengthens the unpublished work of Valls (1978),
who concluded that variation in caryopsis shape was too great to support
the recognition of Diplachne. In particular, the data herein have revealed
nearly continuous variation in cross-sectional shape, ranging from dorsal
compression through circular to various degrees of lateral compression (Table
1; Fig. 2). The lateral/dorsal compression dichotomy of Parodi (1927) sim-
ply does not exist for Leptochloa. Moreover, a nearly identical range of cross-
sectional shapes can exist in closely related genera, as illustrated in a recent
revision of Australian Eragrostis (Lazarides 1997: 176).

The most general observation to emerge from this and other detailed anatomical
and morphological studies of grasses (Davila & Clark 1990, Ellis & Linder
1992: Snow 1996; Lazarides 1997) is that broader sampling regimes gen-
erally reveal additional variation not encountered in narrower surveys. Such
variation cannot be ignored. Future studies therefore should seek both breadth
and depth of sampling in order to minimize the chances of incompletely
characterizing variation. Underestimates of variation leads to errors in the
diagnosis of taxon boundaries and in the accuracy of inferring phylogenetic
relationships, the latter being the very basis by which we make and sup-
port our classifications.

ACKNOWLEDGMENTS

Generous funding was provided by the National Geographic Society (NGS
5594-95) and the Missouri Botanical Garden (Andrew W. Mellon Founda-
tion). A grant from the National Science Foundation to the Population and
Evolutionary Biology Program at Washington University in support of graduate
training is also gratefully acknowledged. My thanks to Dr. John McNeill
and an anonymous reviewer for helping to clarify the manuscript.

280 SipA 18(1)
APPENDIX 1

Voucher bee and herbarium of origin (acronyms follow Holmgren et al. 1990); those lack-
ing herbariu used at MO. For new combinations and new species in Leprochloa see
Snow 1998a, b ane Snow and ane 1997.

y pt fy},

19124 (U S): Snow 6623
L. candata (K. Schum.) N w: Van Someren AH9575 (US); Snowden 1429 (US); Bogdan 130 (UC).
L. chinensis (L.) Nees: aa et ye 6980; Kasim 254 (K). Poore 440 (IK); Davidse 7471 (K,MO); Clayton

5644 (IX).

L. chloridiformis (H 7. ex Stuck.) Parodi: Birabén 179 (LP); Silvews 622 (MICH), Pedersen 3471 (US);

Pedersen 2062 (US).

L. coerulescens a Adam 17175; Adam 14030; Adam 5094,
. canes (R. Boys eae ex Maiden subsp. asthenes (Roem. & Schult.) N. Snow: Snow & Simon 7272;

Snow & Simon 7327; Snow & Simon 7335: Cy risp et al, 2710 (MEL); Thompson & Sharpe HUGI15

(BRI); Fibhand 1008 Ck ).
. decipiens shee beieas Snow et al. 7247; Snow & Simon 7328: Snow & Simon peas — 22548
(CANB); Roe s.n. (MEL, accession 234696); Lazarides an (US); Regan s.n. (CAN
. de i subsp. pre averel en & Betche) N. Snow: Snow & Simon 7323; Snow ae 7329; Snow
Simon 7330; Snow & Simon 7336; Purdie 315D (BRI); Boorman s.n. (G, accession 8227-86) ); Johnson
NB).

713 (CA

& Merr.: Hitchcock 7004 (US); Pringle 6664 (US); Soderstrom 650 (US); MeVaugh

th

t

>

~
\

. digitata (R. Br.) Domin: Snow et al. 7224; Snow et al. 7235; Snow et al. 7246, Burbidge 5326 (CANB),
Blake 11506 (CANB), Blake 6320 (CANB); Walter & Walter 2590 (B).

: si ie S. T. Blake: ied et al. 7228; Snow et al, 7233; Snow et al. 7236: Snow et al, 7241:

Lloyd 979 (CANB); Blake 774

L. sae (Kunth 1) Nees: Snow 5865: one 46783 (NCU); Kral 51801; Mearns 1213 (US); Herndndez

lathus N-2066 (GH); Gould 12183 (K); Castillon 43560 (GH).

Ey en (Nees) T. A. Cope & N. Snow: Snow et al. 6941; Snow & Burgoyne 6954; Snow & Bur ‘goyne

6963; Snow et al. 6982; Schweicker 7 1896 (PRE); Guy & Ward 7 (PRE); Drege s.n. (S, accession 93/
194); mae ve 16419 (P RE).

L, fusca (L.) K subsp. fusca: Snow et al. 7215; Snow et al. 7216; Snow et al. 7222; Snow et al. 7232
Snow et al. rea al. 7237, Tracy 9297 (GH); Fry s.n. (GH).

L. fusca subsp. fascicularts (Lam.) N. Snow: Snow 5786A; Snow 5800; Snow 5804; Snow 5809H: Snow
S811A,; Snow & Koster 5824; Snow & Koster 5840; Snow 5841; Snow 5842; Snow 5896: Snow 5900;
Snow 5901-B; Snow S9O1-E; Snow 5903: Fraser 631 (MICH): Pals 9595 (MEXU); Swatlen 4265
(BAA); Wooton s.n. (BAA

L. fusca cae muelleri ees ) N. Snow: Latz 11 (BRD; Maconochie 13433 (BRI); Must 487 (CANB).

L. fusca subsp. wninervia (J. Presl) N. Snow: Snow & Prinzie 6484; Snow & Prinzie 6567; Snow & Prinzie

6568; Snow 6598.

L. gigantea (Launert) T. A. Cope & N. Snow: Smith 4126 (PRE); Smith 1387 (BRI).

L. ligulata Lazarides: Snow & Simon 7324; Snow 7402; Story & Yapp 25 (CANB).

L. longa Griseb.: se 2612; Soderstrom 1073 (US): Hitchcock . 77 (US).

L. malayana (C. E. Hubb.) Jansen ex Veldkamp: Veiab 25883 (P

L. marquisensis (F. =f .) P.M. Peterson & Judw.: Perlman 14919, oo 15064,

L. monticola Chase: Holdridge 1955 (US); ee H11874 (US).

L. nealleyi Vasey: Snow ne Snow S805A; Snow 5920; Cory 20298 (GH); Waller & Bauml 3096
(GH); Johnston 4872 (MICH).

L. neesti (Thwaites) Benth. Snr & Simon 737
(MO,US); White 8894 (US

L. obtusiflora Hochst.: Mi/ne- ie Taylor 7297 (B), Greenway 10666

Es is (Retz.) Ohwi subsp. panicea: Reekmans 4982 (B); Ahmad s.n. (B, accesssion 175193-120)

itcheock 19474 (US); Field & Loew 6B (US); Backer 35094 (US).

L. fies subsp. brachiata (Steud.) N. Snow: Snow 5905; Snow 5910A; Snow & Prinzie 6529C: Snow &
rinzte 0558; Snow 6654; Snow 6672-A.

L. panicea subsp. mucronata (Michx.) Nowack: Snow 5847A

tap

2

uo

78; Snow & Simon 7384, Davidse & Sumithraarachchi 9180

~~

SNow, Caryopsis morphology of Leptochloa 281

L. panicoides (J. se Hitche.: Snow 5792A; Snow S810A; Snow 6622; Jacob 511 (NCU), Francoer &
Williams ue (ENCB); McDaniel & Rimachi 18020.

L. rupestris Hubb.: Wood 2848 (BM); Wood s.n. (X); Gilbert & Phillips 8874 (K).
L. scabra ee vu 57 fe Snow a Snow S810A; McKenzie & Urbatsch 160 (NY); Nee 31921
Y); Burger é om ay (N

L. southwoodti N. Snow & B. K. . Snow & Simon 7350; Snow & Simon 7362.

L. squarrosa Pilg.: ee 2764 (P); Schlieben 6940 (P); Milne-Redhead & Taylor 7301 (B).

iB a ine N. Snow: Davidse & Sumithraarachchi 9066 (K), Fosberg et al. 50835 (CANB); Clayton
AES).

L. 7, (Backer) Jansen ex Veldkamp: Potlane 15394 (P); Smitinand 3418 (CANB); Kerr 20633 (RK).

L. uniflora Hochst. in A. Rich.: Trimens 28 (US); Rains 67 (US); Thollon 4093 (US)

L. virgata (L.) P. Beauvy.: Pohl & Davidse 10603 (UC); Burkart 22139 (SI); Meza 12; Swallen 3109 (US);
Brickmann 18660 (BAA).

L. eg (Scribn.) Beal: Svow 6597; Ruéz s.n. (ENCB); Palmer 1789 (GH); Goodding & Lusher 139-45

—

(NY
ibs eee P. M. Peterson & Judw.: Hallé 2098 (P).
Cypholepis yemencicus (Schweinf.) Chiov.: Bally & Melville 15783; Verdcourt 3275.

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VAN DEN ae A. and L. ae aa On the classification of the Chloridoideae (Poaceae).
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XANTHOSTEMON FRUTICOSUS (MYRTACEAE), A
NEW SPECIES FROM THE PHILIPPINES

PETER G. WILSON
National Herbarinm of New South Wales
Royal Botanic Gardens
Sydney, N.S.W. 2000, AUSTRALIA
LEONARDO L. CO
Herbarium, Institute of Biology
University of the Philippines
Diliman 1101, Quezon City, PHILIPPINES
ABSTRACT
A new species of Xanthostemon, X. fruticosus, presently known only from the Palanan Wilderness
in Isabela Province, Luzon, is described and its relationships discussed.
RESUMEN

Una nueva especie de Xanthostemon, X. fruticosus, en la actualidad solamente cope icae: de
la Palanan Wilderness en la provincia de Isabela es descrita y sus rel

discutidas.

Xanthostemon is a genus of around 45 species that occurs in Australia, New
Caledonia, the Solomon Islands, Papua New Guinea, Indonesia (Irian Jaya,
the Moluccas and Sulawesi) and the Philippines. In the Philippines, only
four species have been named and there has been little published on the
taxonomy of the genus since Merrill (1952) clarified the application of the
names X. speciosus and X. verdugonianus.

In 1991, a preliminary inventory of the Palanan Wilderness was under-
taken as a joint project of Conservation International, Leiden University
and the Isabela State University. A report of this inventory is given by Co
and Tan (1992). One of the vegetation types encountered was a low scrub
that occurred on ultrabasic substrate and was dominated by a shrubby
Xanthostemon species with large, right red flowers. This has proven to be an
undescribed species, most closely related to X. verdugonianus and is here described
as X. fruticosus.

Xanthostemon fruticosus Peter G. Wilson & L. Co, sp. nov. (Fig. 1). Tyre:
PHILIPPINES. Luzon Istanp. Isabela Prov.: Aubarede Peninsula, Lanay acne ca.
17 km NNW of Palanan Point, 25 May 1991, aie Co 3583 (HOLOTYPE: A; ISOTYP
PNH, PUH, CAHUP, ISB, L, KEP, US, K, CANB).

Sipa 18(1): 283-286. 1998

284 Sipa 18(1)

Lesle
Bad,

Fic. 1. Xanthostemon fruticosus Peter G. Wilson & L. Co. A. Habit B. Immature fruit show-
ing neatly circumscissile hypanthium. C. Open, mature fruit. D. Detail of leaf showing
venation. All from holotype. Scale bar: A, D=3 cm; B, C=1 cm.

WILSON AND Co, A new Xanthostemon species 285

Xanthostemon verdugiano hypanthio circumscisso valde affinis sed habitu fruticoso foliis
floribusque maioribus pedicellis pedunculisque longioribus et petalis roseis differt.

Shrub ca. 1 m high; young shoots sparsely appressed-hairy, glabrescent.
Leaves spirally orale petiole 7-10 mm long, 2.5—3 mm wide; lamina
obovate, 6—8 cm long, 3.5—4 cm wide, length:breadth ratio 1.6—2.1:1, co-
riaceous, apex rounded and emarginate; oil glands numerous. Inflorescences
crowded toward the apex of seasonal growth unit, subtended by slightly
reduced foliage leaves, axillary, the apical bud apparently always growing
on (i.e. shoot auxotelic). Unit inflorescences triads; peduncles 5-9 mm long,
the upper ones shorter giving a corymb-like appearance to the flowering
shoot; pedicels 6.5—10 mm long. Hypanthium cup-shaped, 5—6 mm in diameter,
4—5 mm deep, exceeding the summit of the ovary, + glabrous. Petals 5, pink,
oblong to obovate, 5.5—8 mm long, 3.5—6.8 mm wide. Sepals 5, equal, 2—2.7
mm long, 2.5—3.8 mm wide, +triangular, apex obtuse to rounded. Stamens
red +equally spaced, ca. 22 ina single series around the rim of the hypanthium,
filaments 22—25 mm long; anthers 1.2—1.5 mm long, connective broad with
one large gland at the apex and 2—5 other, smaller glands. Ovary partly in-
ferior, (2—)3-locular, glabrous. Style 30-35 mm long, extending 3-5 mm
beyond the stamens; stigma as wide as or slightly narrower than the style,
flat. Fruit depressed-globular, ca. 8 mm diameter, ca. 6.5 mm high, broadly
lobed at each loculus; hypanthium neatly circumscissile 1.5—2 mm above
the base, the remnant +flattened under the capsule. Placentas rod-like, horizontal.
Seeds not seen.

Distribution.—Known only from the type locality.

Ecology.—Xanthostemon fruticosus has been found near the coast in a low
scrub community occuring on exposed sites on soils derived from an ultra-
basic substrate. It is the dominant component of this community type at
Lanay which also includes a number of other genera of spreadiing or erect
shrubs (Co & Tan 1992).

Conservation status. —Apparently rare. Only recorded from one of four ultrabasic
sites visited in this part of north-eastern Luzon. Ultrabasic areas are very
marginal for agriculture but are potentially subject to applications for nickel
mining. As a wilderness, the region does have legal status as a ‘Protected
Area’ but this may not be an impediment to mining.

Etymology.—The epithet is derived from the Latin, fruticosus, shrubby, referring
to the distinctive habit, a characteristic that distinguishes it from the re-
lated X. verdugonianus.

The red-flowered X. speciosus Merr. From Palawan and nearby islands has
a broader, dish-shaped hypanthium with distinct vesicles and is thus not
closely related to the new species but belongs with X. confertiflorus (Sulawesi)
and X. youngii (north-eastern Australia) in sect. Vestcaria of Gugerli (1940).

286 Stipa 18(1)

Xanthostemon speciosus is usually a tree of 5 meters or more but small trees or
shrubs of only 1 meter or so in height have been recorded (e.g. Co 3032, A,
P , KEP)

Xanthostemon fruticosus is closely related to X. verdugonianus which it re-
sembles in having red flowers with a circumscissile hypanthium. Xanthostemon
verdugonianus is a tree up to 26 m high that ranges from north-eastern Mindanao
to Sibuyan Island, the latter being over 500 km south of the Palanan area.
Despite its range, X. verdugontanus has a very limited habitat and is much
sought-after for its durable timber, making it a threatened species (Yao &
Ulep: 1981, 1985).

The neatly circumscissile hypanthium appears to be unique to this pair
of species. Two species in New Caledonia have hypanthia that split and are
irregularly circumscissile but these are members of an endemic group of
taxa with bullate leaves (sect. Bud/ata of Gugerli 1940) and do not appear
to be closely related to the Philippine species.

The species may be distinguished as follows:

1. Shrub ca. 1 m tall; lamina 6-8 cm long, 3.5—4 cm wide; petiole 7-10 mm
long; petals pink. X. fruticosus
1. Tree to 26 m tall; lamina 3-7 cm long, 1—3.5 mm; petiole 4-7 mm long;

petals red. X. verdugonianus

ACKNOWLEDGMENTS

The first author is grateful to the directors of A and CANB for loans of, or
access to, herbarium specimens examined for this paper. Thanks, also, to
Lesley Elkan for the illustration and Jaime Plaza for assistance with the Spanish
abstract. The second author thanks Conservation International for funding
his field work in the Palanan Wilderness in 1991.

REFERENCES
Co, L.L. and B.C. Tan. 1992. Botanical exploration in Palanan Wilderness, Isabela Prov-
ince, the Philippines: first report. Fl. Males. Bull. 11:49—53.
GuceERLI, K. 1940. era der Myrtaceengattung Xanthostemon. Repert. Spec. Nov.
egni Veg. Beih. 120:1-149.
Merritt, E.D. 1952. Notes on <a F Muel
Arbor. 33:150-165.
Yao, C.E. and E.V. Utep. 1981, Mangkono in Babatngon. Canopy 7:5.
Yao, C.E. and E.V. Utsep. 1983. More on Mangkono. Canopy 9:6—7.

=

er and Kje//bergiodendyon Burret. J. Arnold

SEASONAL CHANGES IN CONCENTRATION
AND DISTRIBUTION OF HEAVY METALS
IN CREOSOTEBUSH, LARREA TRIDENTATA
(ZYGOPHYLLACEAE), TISSUES IN THE EL PASO,
TX/CIUDAD JUAREZ, MEXICO AREA

WILLIAM P. MACKAY, RICHARD MENA, NICHOLAS E.
PINGITORE JR., KEITH REDETZKE, C. EDWARD FREEMAN,
HAROLD NEWMAN, JOHN GARDEA, and HECTOR NAVARRO

Laboratory for Environmental Biology and
Department of Biological Sciences (WPM, RM, CEF, HN, JG, HN, KR)
Department of Geological Sciences (NEP)
University of Texas, El Paso, TX 79968, U.S.A.

ABSTRACT

We compared seasonal changes in concentrations on a elements, arsenic, cadmium,
copper and lead, in samples of various tissu -reosotebush collected in the El Paso, USA/

Ciudad Juarez, Mexico region during 1 980- 81 “ed 1994— 95. Levels in leaf tissue need
seasonally, with concentrations dropping in the spring and late fall, corresponding to the
time of leaf drop in the plant. This suggests that most of the heavy metals were simply
deposited on the surface of leaves, although data are presented which indicate that internal
tissues also have significant levels of heavy metals. These seasonal cycles were less pronounced
in the most recent samples. Levels of all four elements in cresotebush were significantly
high in a region, as compared to a distant control area. Concentrations of most elements
dropped below detection limits within 12 to 30 km from areas of highest concentration.
Densities of native Chihuahuan Desert flora and lichens are low in the area, suggesting an
apparent negative impact of industrial pollution on the local vegetation. Cadmium and
lead levels in creosotebush tissues have dropped over the past 15 years, suggesting that
enforcement and strengthening of environmental laws has reduced the air pollution levels
in the El Paso area.

RESUMEN

Comparamos los cambios estacionales de las concentraciones de 4 elementos, arsénico,
cadmio, cobre y plomo, en muestras de varios tejidos de gobernadora colectados en el area
de El Paso y Ciudad Judrez, México durante 1980-81 y 1994—95. Se encontro un cambio
estacional de los metales con las concentraciones disminuyendo en la primavera y otofo,
que coresponden al momento de la caida de las hojas de la planta. Esto sugiere que hay
deposicién de los elementos en la superficie de las hojas, aunque presentamos datos de que
hay acumulacién de metales en los tejidos interiores. Estos ciclos estacionales estan menos

pronunciados en los tltimos afios. Los niveles de los elementos son altos en la region, pero
estan concentrados en un 4rea entre 12 y- 30 kil6metros del drea con concentraciones mas
altas. Las densidades de la flora y liq | drea son bajas

de la polucion industrial en la vegetaci6n local. Los niveles de cadmio y eee en los tej ne

Sipa 18(1): 287-296. 1998

288 Sipa 18(1)

ee | | ad iG ]

ou

ha disminuido en los tltimos 15 afios, log r | |
ha reducido los niveles de polucién en el area de El Paso.

The El Paso/Cd. Juarez area is one of the many regions along the United
States/Mexico border which suffers from environmental contamination. The
area has been subjected to heavy industrial activity, including smelting and
hydrocarbon refining, for over 100 years. This has resulted in an accumula-
tion of heavy metals, especially lead, cadmium, copper and zinc in the local
soils (Barnes 1993; Ndame 1993). Arsenic contamination has been previ-
ously documented to occur in this region of west Texas (Shields 1991).

These elements and their compounds can cause medical problems in humans
and other animals (Elbahri & Benromdane 1991; Louekari et al. 1991). Negative
effects of heavy metals on plants have been documented (D’itri 1982; Fernandes
& Henriques 1991), including the prevention of the uptake of potassium
and phosphorous by roots. Moreover, copper may damage chlorophyll and
increase the potency of fungal diseases (Connell & Miller 1984). These ef-
fects may eliminate some plant species, with concomitant increases in the
abundances of others, thus changing plant community structure. Plants may
tolerate heavy metal contamination (Connell & Miller 1984) by excluding
metals from sensitive tissues, modifying metabolic pathways to prevent damage
or assembling specific enzymes to detoxify heavy metals. The specific ef-
fects of heavy metals on the flora of the northern Chihuahuan Desert have
not been documented, but Worthington (1989) previously reported a re-
duction in species richness in native plants in this area, presumably the result
of such contamination.

In this investigation, tissues of the dominant Chihuahuan Desert shrub,
Larrea tridentata were analyzed, to document continued heavy metal con-
tamination in western Texas. This work is part of a larger study on the ef-
fects of heavy metal contamination on the flora and fauna of the northern
Chihuahuan Desert and the investigation of bioaccumulation into higher
trophic levels.

MATERIALS AND METHODS

We selected a site on the campus of the University of Texas (UTEP) in
western El Paso to follow seasonal change in heavy metal concentrations in
creosotebush. The site is located within 2 km ofa local smelter, which ap-
pears to be a point source for the high concentrations of heavy metals in the
area. Four elements were included in the analysis: Arsenic, Cadmium, Copper
and Lead. Additional sites used to determine the distribution of heavy metals
in creosotebush tissues in the region included 62 sampling areas near the
border in the United States and six in Mexico within 10 km of the border
and of the smelter, of which 52 yielded creosotebush specimens (five repli-

MAcKay, ET AL., Heavy metals in Larrea tridentata 289

cate bushes at each site). Sites were selected which had native Chihuahuan
Desert vegetation located at least 50 meters away from any road. Prelimi-
nary analyses showed higher levels of lead next to roadways, which decreased
to background levels within a few meters, an effect also noted by Motto et
al. (1970), Gratani et al. (1992) and Lebreton and Thevenot (1992).

Leaves were stripped, and twigs were cut from the plants. Roots and trunk
tissues were also harvested from plants. Tissues were placed in numbered
paper bags and returned to the laboratory. The samples were further cleaned,
removing all foreign matter, but were not washed. Bark and exterior tis-
sues were carefully removed from pieces of trunks, to avoid contamination
of internal tissues. Tissues from 1980-1981 were collected North of the
Education Building on the UTEP campus, as part of another study (Free-
man 1982). The tissues were ground and stored dry in glass vials until they
were analyzed in 1994. Tissues from 1994-1995 were collected through-
out the season from five specific bushes, located in Charlie Davis Park, on
the campus of UTEP. This allowed a comparison of the percentages of heavy
metals in the tissues of each of the five creosote bushes.

Approximately 20 grams of tissue were placed in a crucible and muffled
for three days at 425°C. This relatively low temperature was selected to
avoid vaporizing the four elements or their salts (based on the recommen-
dations of Dr. Jim Rayon and Dr. Ken Dodson of the Environmental Pro-
tection Agency). After ashing, 100 mg of ash were dissolved in 20 ml of
14% nitric acid (V:V), without filtering. Reagent grade acid and double
glass distilled water were used for all solutions. Samples were prepared and
stored in glass scintillation vials with polypropylene-lined caps

Samples were analyzed on a Beckman SpectraSpan 6 direct current plasma
atomic emission spectrograph (DCP-AES). We followed the protocol of EPA
method 6010 for the inductively coupled plasma emission spectrometer (US
EPA 1986). Three of the wavelengths used are those recommended by EPA:
193.696 nm for arsenic, 213.598 nm for copper, and 220.353 nm for lead.
The fourth wavelength, 228.802 nm, was substituted for the cadmium analysis
to achieve the lowest detection limit. The DCP was calibrated with pre-
pared standard solutions such that the linear calibration curve had an R? o
at least 0.995. The samples were analyzed by DCP three times and reported
the mean and standard deviation of these readings. If the relative standard
deviation was greater than 3%, the sample was reanalyzed. Blanks, dupli-
cate samples and prepared standards (spiked samples) were analyzed at least

—

once every ten samples for quality assurance/quality control. Cadmium
concentrations in the samples collected in 1994-1995 were close to or be-
low the limits of detection and therefore we have not presented them. Af-
ter analysis, the DCP solution values were converted to ppm in the original

290 Sipa 18(1)

ash. The detection limits in the ash for this study were 200 parts per mil-
lion (ppm) for arsenic, 11 ppm for cadmium, 85 ppm for copper and 130
ppm for lead.

Data were plotted with SURFER (Golden Software Inc., Golden Colo-
rado), using the grid method with inverse squared distances and the sur-
face module (Keckler 1995).

RESULTS AND DISCUSSION

Seasonal changes.

Levels of all four elements varied seasonally during both
time periods (Fig. 1). Levels were high in the winter and showed a small
drop in the spring when there is a combination of minor leaf drop and strong

spring winds. The levels increased during the summer, especially during
the early sampling period (Figs. la, b,c), but later showed a large drop during
the fall when creosote lose a large proportion of their leaves (Mackay et al.
1987). Concentrations increased again during the winter months, especially
during 1980-1981. Levels of heavy metals were lower in the 1994-1995
samples (Fig. 1d & f). Cadmium and lead were both approximately 4 times
higher in 1980-1981, suggesting that enforcement and strengthening of
environmental laws has reduced the air pollution levels in the El Paso area.
Seasonal changes in concentrations of all metals during both sampling
periods were statistically significant (Table 1). The first harmonics from Fourier
analysis of the means (Little & Hills 1972), corresponding to the seasonal
effect, were significant for all years, alchough the patterns were more pro-
nounced during the first sampling period (Fig. 1) and had correspondingly
higher F values (Table 1). Other harmonics were statistically significant in
1980-1981, showing the importance of leaf drop in lowering heavy metal
concentrations during these years, alchough there was no pattern in which
harmonic was significant after the first. The lack of significance of higher
harmonics during the second sampling period suggests that leaf drop was
not significant in reducing heavy metal content during those years.
Contamination of tissues. —It appears that large amounts of the heavy metals
arsenic, copper and lead are deposited on the leaves, but heavy metals are
also incorporated into the tissues of this plant (Fig. 2). Large concentra-
tions of all three elements were found in the roots. The trunks also had high
concentrations of the three elements, which were present in the internal
tissues as well. The branches and the leaves also had high concentrations of
the three elements. These data suggest that heavy metals may be deposited
on the outside of the plant, but is also taken up by the roots and incorpo-
rated into the woody tissue
Heavy metal geographical distribution: Levels of the four elements (ar-
senic, cadmium, copper and lead) in creosotebush ash were elevated adja-
cent to the United States/Mexico border on the west side of El Paso (Fig.

Mackay, ET AL., Heavy metals in Larrea tridentata 29]
500 500
400 — “ 1980 - 198] 400 + 1994 - 1995
300 + 300 +
200 - 200
100 - 100 -
04 4 d
AS a) ° AS )
Pos 2 le lanl coral a oa aa ee Foor Fa
180 6000
MT 160 - 1980 - 1981 1994 - 1995
— 140 5 5000 4
120 -
4000 -
< sel
= eal 3000 +
40 5 2000 +
PX 204¢4 b) Cu e)
0 fs oe a ae 1000 [oon eee IS
ene 1400
1980 - 1981 | 1994 - 1995
5000 - 1200 -
4000 4 1000 -
3000 800
600 -
2000 oe
1000 - 4
» ee c) Pb f)

ep gens nee aia
A OD F A
Fic. 1: Seasonal changes in heavy metal concentrations in the leaves of the creosotebush,

Larrea tridentata during 1980-1981 and 1994-1995. The gap in 1994-1995 is due to

lost samples. Error bars represent standard error of the mean, and are not included ina, b,

and ¢ as many of the data points are based on a single sample. The months are abbreviated
by a single letter on the x axis.

3). Levels of cadmium were as high as 190 ppm in ash, those of copper reached
5200 ppm and lead levels were as high as 1200 ppm (Fig. 3, note that the
surfaces in the figures are close to average values and do not extend to these
extreme values). The highest levels of the three elements were recorded on
the east side of Mount Cristo Rey (Peak 1 in Fig. 3a), on a small mesa west
of McNutt Road (Peak 2 in Fig. 3a), and on the UTEP campus (Peak 3 in
Fig. 3a). Levels of all three elements were lower on the southwest side of
Mount Cristo Rey, perhaps due to a wind-shadow effect. Levels of all three
elements rapidly decreased to the east and fell below detection limits at
distances of between 12 and 30 km (Fig. 3).

292 Sipa 18(1)

Tate |. Results of Fourier Analysis (Little & Hills 1972) of the periodic functions in Figure 1. Means

were used in the analysis.

Source df Mean Squares F Values

Arsenic, 1980-1981

First harmonic 2 108121.6 22.2°™°
12 4880

Second harmonic 2 9086.0 2.2 ns
error 10 4039.4

Third harmonic 2 10400.7 4.2"
error 8 2449.1

Fourth harmonic 2 5426.9 3.7 ns
error 6 1456.5

Fifth harmonic Z 1361.7 0.9 ns

4 1503.8
Cadmium, 1980-1981

First harmonic 2 8798.9 9.4"
error 12 940.7

Second harmonic 2 2910.9 5.3"
error LO 546.7

Third harmonic 2 28.6 O ns
error 8 676.3

Fourth harmonic 2 LOSL.O 1.9 ns
error 6 551.1

Fifth harmonic 2 1627.3 125.50"

4 3.0
Lead, 1980-1981

First harmonic 2 12113720.8 19.9"
error 12 608560.6

Second harmonic 2 1801911.0 4.9"
error 10 369890.5

Third harmonic 2 292706.7 0.8 ns
error 8 389186.5

Fourth harmonic 2 423497.9 L.1l ns
error 6 377749.3

Fifth harmonic Z 732260.3 3.7 ns
error { 200493.9

Arsenic, 1994-1995

Firse harmonic 2 198 26.3 5.9
error 12 3362.4

Second harmonic 2 1159.2 0.3 ns
error 10 3991.7

Third harmonic 2 3697.3 L.7 ns
error 8 3309.7

Fourth harmonic 2 1123.4 0.2 ns
error 6 4733.9

Fifth harmonic | 2 6111.7 3.1 ns
error 4 1978.4

Mackay, ET AL., Heavy metals in Larrea tridentata 293

TABLE 1. continued.
Source df Mean Squares F Values
Arsenic, 1980-1981
Copper, 1994-1995

First harmonic 2 1168398.7 5:07
error 12 232963.0

Second harmonic 2 352562.1 1.8 ns
error 10 198791.8

Third harmonic 2 231261.4 1.2 ns
error 8 185804.0

Fourth harmonic 2 118186.9 0.5 ns
error 6 230882.1

Fifth harmonic 2 267145.6 1.7 ns
e 4 158355.2

Lead, 1994-1995

First harmonic 2 68884.3 7a
error 12 9648.9

Second harmonic 2 8773.5 0.9 ns
error 10 9899.0

Third harmonic 2 8648.6 0.8 ns
error 8 10399.2

Fourth harmonic 2 2035.2 0.1 ns
error 6 15975.2

Fifth harmonic 2 15531.3 0.9 ns
> 4 16863.0

= F value of harmonic significant at 0.05 level, ** = F significant at 0.01 level, *** = F signifi-

cant at 0.001 level, ns = F value not significant.

The areas with the highest values of As, Cu, Cd, and Pb in creosotebush
(peaks 1—3, Fig. 3) coincide with the sites of the highest concentrations of
heavy metals in the soils (Barnes 1993; Ndame 1993) and in fluff grass (MacKay
et al. 1998). In this locale, Pb in the soils exceeds the EPA TCLP (Toxicity
Characteristics Leaching Procedures) regulatory limit at a number of sites,
and Cd is reported quite close to the limit. The spatial correspondence between
elevated metal levels in the soil and the flora is not unexpected. The mechanism
of uptake of heavy metals by creosotebush remains to be elucidated.

Densities of native Chihuahuan Desert flora and lichens are low in the
area (Worthington 1989, MacKay et al. in prep.), suggesting an apparent
negative impact of industrial pollution on the local vegetation. The spe-
cific effects of heavy metals on the flora of the northern Chihuahuan Desert
are currently being documented. The demonstrated reduction in species richness
in native plants in this area is presumably the result of heavy metal con-
tamination from the smelter. These effects may eliminate some species and
increase the abundance of others.

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Fic. 2: The distributions of lead, arsenic and copper in specific tissues of the cre

from Charlie Davis park on the University of Texas, El Paso campus, E] Paso, ”

ves
Sr
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ACKNOWLEDGMENTS

We would like to thank the United States Department of Agriculture,
especially J.A. Vigil and J. Underwood, for permission to import plant tis-
sues from Mexico. The Texas Department of Parks and Wildlife, especially
David Riskind, allowed us to collect plant tissues in the Franklin Moun-
tains State Park. Ken Dodson and Jim Rayon offered suggestions regard-

ing sample preparation. Our research was supported by the Center for

InterAmerican and Border Studies of the University of Texas and by the

National Science Foundation (HRD 9253021).
REFERENCES

Barnes, B. 1993. An evaluation of metals concentrations in surficial soils, El Paso County,

Texas. Unpublished Master's Thesis, University of Texas at El Paso.

MAckay, ET AL., Heavy metals in Larrea tridentata 295

3
Pee DISTRIBUTION OF ARSENIC a3 DISTRIBUTION OF CADMIUM
Ba. 3
g x'
= g4
d ME ay

33
Rs x
ed & 8.
a a
a

PPM COPPER IN ASH
Bar t0 {SS
i

6er.e

oy] |)

Fic. 3: The distribution of arsenic, cadmium, copper and lead in the E] Paso/Ciudad Juarez
area. Charley Davis Park on the UTEP campus is located at the origin in the x-y coordi-
nate system (0,0). Peak one corresponds to the east side of Mount Cristo Rey, Peak 2 toa
mesa above (west of) McNutt Road and Peak 3 is on the UTEP campus. Detection limits
were 200 ppm for arsenic, 11.2 ppm for cadmium, 84 ppm for copper and 132 ppm for

ead.

ConneELL, D.W. and G.J. MILLER. 1984. Chemistry and ecotoxicology of pollution. John
Wiley and Sons, New York.
D’irri, EM. 1982. Acid precipitation: effects on ecological systems. Ann Arbor Science,
Ann Arbor, MI
ELBauri, L. and S. BENROMDANE. 1991. Arsenic poisoning in livestock. Veterinary Human
Toxicol. 33:259-26
FER RNANDES, J.C. and a HE Es. 1991. Biochemical, eer and structural ef-
cts of excess copper in plants. Bot. Rev. 57:246-262
ae C.E. 1982. Seasonal variation in leaf nitrogen in rere: (Larvea tridentata
DC.} Cov.: ra ieee SouthW. Naturalist 27:354—-3
GraTANI, L., 8. TAGLIONI, and M.F. Crescente. 1992. The cman of lead in agricul-
tural soil ond vegetation mies a eee C hemosphere 24:941—94¢
Kecker, D. 1995. Surfer for Windows, Ver 6.01, Golden aan Inc., Golden Colo-
rado
LeBRETON, L. and D.R. THEvenor. 1992 Sees metallique relargable par les aerosols
d'origine autoroutiere. Environ. Technol. 13:35—44.
Lirtte, T.M. and F. Hints. 1972. Re ae in agricultural research. Agric. Ext.
Service, Univ. California.

296 Stipa 18(1)

LoueKARI, K., $. VALKONEN, S. Pousi, and L. VirTANEN. 1991. Estimated dietary ita of
lead and cadmium and their concentration in blood. Sci. Total Environ. 105:87—99.
Mackay, W.P., S. Sitva, S. Lorinc, and W. Wuirrorp. 1987. The role of subterranean ter-
mites in the decomposition of above ground creosotebush litter. Sociobiology 13:235—

2D.

Mackay, W.P., L. De La Rosa, M. Zavata, P. SHELTON, J. GARDEA, R. MENA, N. PINGITORE,
and G Crawrorb. 1998. The distribution of heavy metal contamination in the El Paso,
TX/Ciudad Juarez, Mexico area, using fluff grass, Erionewron pulchellum, as an indicator
organism. Proc. Chih. Desert Res. ie (In Pres

Morro H.L., R.H. Daines, D.M. CuitKo, and C.K. ae TO. 1970. Lead in soils and eae
its relationship to traffic une and proximity to highways. Environ. Sci. Technol. 4:23 1—

2

Nbame, E.C. 1993. Heavy metals in soils in the vicinity of the University of Texas at El
Paso campus (El Paso, Texas). Unpubl. Master’s Thesis, University of Texas, El Paso.

SHiELps, J. 1991. Ambient air arsenic levels along the Texas - Mexico border. J. Air Waste
Manag. Assoc. 41:827—

US ENVIRONMENTAL PROTECTION AGENCY. 1986. Test methods for evaluating solid waste
(third edition). Washington, D.C.

WORTHINGTON, R.D. 1989. ae of El Paso pollutants on the lichen, moss, and winter
annual flora on andesite rock formations. In: P. Ganster and H. Walter, eds. Environ-

fon

mental hazards and bioresource management in the United States/Mexico borderlands,
SLA Latin American Center Publications. Los Angeles. Pp. 105-115.

NEW PLANT RECORDS FOR DOMINICA,
LESSER ANTILLES

STEVEN K. HILL
Center for Biodiversity
Illinois Natural History Survey
607 E. Peabody Drive
Champaign, IL 61820, U.S.A.
ARLINGTON JAMES
Division of Forestry and Wildlife
Botanic Gardens
Rosean, COMMONWEALTH OF DOMINICA
ABSTRACT

Twenty-six vascular plant species are reported here for the flora of Dominica, in the Lesser
Antilles. These include two pteridophytes, 13 monocotyledons, and 11 dicotyledons. Four

of these species are reported as new to the Lesser Antilles: Marsilea polycarpa Hook. & Grev.,
Diplacrum capitatum (Willd.) Boeck., Rhynchospora racemosa Wright ex Sauvalle, and Pharus
lappulaceus Aubl.

RESUMEN
Se citan 26 plantas vasculares adicionales para la flora de Dominica en las Antillas Menores.
Estas especies incluyen dos pteridéfitos, 13 monocotiledéneas y 11 dicotiled6éneas. Cuatro
de estas especies se presentan como nuevas para la flora de las Antillas Menores: Marsilea
polycarpa Hook. & Grev., Diplacrum capitatum Willd.) Boeck., Rhynchospora racemosa Wright
ex Sauvalle, y Pharus lappulaceus Aubl.

INTRODUCTION

Dominica is a volcanic island located between the French islands of Guadeloupe
and Martinique in the Lesser Antilles. It is about 45 km long and 24 km
wide. While it is a small island (1088 sq. km), it presents a diversity of
habitats along altitudinal and moisture gradients because of its rugged
mountainous topography. Several of the mountain peaks exceed 1200 m
(Morne Diablotins 1433 m, Morne Trois Pitons 1402 m, Morne Watt 1242
m, Morne Anglais 1223 m). Water is abundant on the upper slopes of the
mountains (e.g. at Freshwater Lake, 8459 mm of rain/yr. has been recorded)
but rainfall on the western (Caribbean) coast is severely restricted by a rain
shadow (< 2000 mm/yr.) and a dry scrub forest prevails. Severe damage caused
by hurricanes and agriculture has also provided opportunities for opportu-
nistic species, increasing the diversity. The coastline is very steep, and the

Sipa 18(1): 297-305. 1998

298 Sipa 18(1)

clifts generally continue precipitously into the ocean. Level land is prima-
rily restricted to river deltas in a few narrow bands near the coast, where
most citizens live.

Botanically, Dominica is rather well-known. The dicotyledonous flora
was treated by Nicolson (1991) who estimated a vascular flora of 1226 spe-
cies. The most recent treatment of its pteridophytes and monocots was Howard's
six volume Flora of the Lesser Antilles (1974, 1977, 1979, 1988, 1989a, 1989b)
which included detailed distribution notes within that region, and treated
the dicots as well. Whitefoord (1989) added 40 phanerogams and six ferns
to the flora. This paper reports twenty-six vascular plant species newly dis-
covered and vouchered for Dominica.

METHODS

The plants reported here as new to Dominica were discovered during the
period 1990-1997, Approximately 2000 collections were made by the first
author for the purpose of assembling a reference herbarium at the Spring-
field Estate, formerly called the Archbold Tropical Research Center after
the late John Archbold who donated the property to Clemson University
in 1989. The Springfield Estate is currently leased from Clemson Univer-
sity by a Dominican non-government organization, the Springfield Centre
for Environmental Protection, Research, and Education (SCEPTRE). The
herbarium there has been informally designated with the acronym “atrc.”
Specimens donated by several other collectors have also been incorporated
into the herbarium.

Collections (2005 numbers) have been made on Dominica during eight
visits to the island by Hill as follows: 20 March—26 March 1990 (collec-
tion numbers 27230-21379), 4 March—23 March 1991 (22009-22179),
26 July—5 August 1992 (23924-24171), 5 March—16 March 1993 (24612—
24847, w/J. Gable & B. Dorsey), 7 December—19 December 1993 (25309—
25563, w/I.Renne & D.Bradshaw), 28 May—6 June 1994 (25610-25872),
24 May—9 June 1996 (27861-28262), and 23 February 1997-14 March
1997 (28868-29109, w/L.R. Phillippe). Sets of specimens have been de-
posited primarily in che Dominican herbarium (atrc), Clemson University
(CLEMS), the Illinois Natural History Survey (ILLS), and the Smithsonian
Institution (US). Additional duplicates have been distributed to herbaria
cited after the individual specimens.

New records of species generally considered to be indigenous to the Lesser
Antilles (according to Howard 1974, 1977, 1979, 1988, 1989a, 1989b)
are reported here. Some may have actually been introduced to Dominica by
human activities, but direct evidence is lacking.

en)

HI AND James, Plant records for Dominica 299
ANNOTATED PLANT LIST
PTERIDOPHYTES

HYMENOPHYLLACEAE
Trichomanes crinitum Sw. An epiphyte similar to Trichomanes crispum L. but
differing in its lobed or pinnatifid, not entire, pinnae. Its general distribu-
tion is northern South America to Jamaica, and it has been reported from
Guadeloupe, St. Vincent, and Grenada in the Lesser Antilles. We thank A.R.
Smith for the identification

St. George Parish: elfin forest on windswept ridge between Breakfast River and Valley
of Desolation, 6 Mar 1997, Hill & Phillippe 29070, (atrc, ILLS, UC).
MARSILEACEAE
Marsilea polycarpa Hook. & Grev. A trailing perennial of temporary ponds,
this species of northern South America has not been previously reported in
the Lesser Antilles. It has several globose sporocarps arranged and attached
in a linear fashion on the frond stipes rather than the single basal sporocarp
seen in most other species of the genus. Neither the genus nor family have
been previously reported from Dominica.

St. Andrew Parish: locally abundant in and around temporary ponds at E end of Melville
Hall Airport, Londonderry Bay, 23 Feb 1997, Hill & Phillippe 28868 (atrc, BRIT, ILLS,
MO, NY, UC, VT)

FLOWERING PLANTS
LILIOPSIDA

AGAVACEAE

A gave dussiana Trel. The century plants are seldom collected, and none were
reported from Dominica by Howard (1979). This species, which is endemic
to the Lesser Antilles, is very conspicuous in the dry scrub on the dry rocky
hills of the NW coast of Dominica in areas of very low rainfall, growing
mostly with cacti. Previous collections are known from St. Barts, Antigua,
Montserrat, ease and ESS aS:

St. Joseph Parish: Morne Raquette ), Au Piton Road, 18 Dec 1993, Hill 25524
(atrc, NY). St. Peter Parish: ‘Ciemen t James’ property, Anse a Liane trail to coast, 0.9 km
N of Colihaut, 2 Mar 1997, Hill & Phillippe 58988 ee BRIT, ILLS, MO, NY, TAES,
USF, VT).

CYPERACEAE

Diplacrum capitatum (Willd.) Boeck. This species is rather common in Ven-
ezuela and ranges from Panama to Bolivia as well. This appears to be the
first record from the Lesser Antilles. Howard (1979) listed only Diplacrum
longifolium (Griseb.) C.B. Clarke ex Dur. & Schinz. for the Lesser Antilles,
rom St. Paul Parish, Dominica. We thank M. Strong for the identification.

300 Sipa 18(1)

St. Joseph Parish: Imperial Highway, Central Forest Reserve, Dleu Gommier area, 6
Jun 1994, Hill 25864 (atrc, NY, US, USF).
Fimbristylis cymosa R. Br. subsp. spathacea (Roth) T. Koyama. This is a char-
acteristic, wiry-leaved and tough-rooted, sedge of the windswept flats of
the E and NE coasts of Dominica. Pantropical in distribution, it has been
collected previously on most of the Lesser Antilles, including Guadeloupe
and Martinique.
St. Andrew Parish: roadside 0.5 mi N of Melville Hall Airport, Londonderry Estate,
16 Mar 1993, Hill 24842 (atrc, CLEMS, MO, NY, TAES, USF).
Rhynchospora racemosa Wright ex Sauvalle. This is the first report of this species
for the Lesser Antilles. It is relatively common in the Greater Antilles. We
thank M. Strong for the identification
t. Peter Parish: Morne Espagnol, from Coastal Highway to summit, 4 Aug 1992, Hi//
24149 (CLEMS, NY).
ORCHIDACEAE
Brachionidium parvum Cogn. This tiny epiphyte of the upper montane rainforests
was previously known from Venezuela and from Guadeloupe, Martinique,
and Grenada in the Lesser Antilles. It was to be expected in Dominica but
probably has been overlooked because of its small size (< 5 cm tall). We
ae D. Nicolson for the identification.

. Peter Parish: NW side of Morne Diablotins peak, elfin forest, 6 Mar 1993, Hi//
ae (atrc, CLEMS). St. George Parish: Laudat, ca. | km W of Titou Gorge, 4 Jun 1996,
Hill 28177 (atrc, ILLS, USF).

POACEAE
Ichnanthus nemorosus (Sw.) Doell. var. swartzi7 K.E.Rogers. Generally distributed
in the Antilles and Trinidad, this trailing rainforest species has been col-
lected previously on St. Kitts, Nevis, Montserrat, Martinique, St. Vincent,
and Grenada in the Lesser Antilles.

St. George Parish: Morne Anglais trail to summit, $ of Giraudel, 2 Jun 1996, H7//
28124 (atrc, ILLS).

lsachne angustifolia Nash. This thicket-forming grass which often climbs over
other plants has been known previously from Puerto Rico, and from Guadeloupe,
Martinique and St. Kitts in the Lesser Antilles. The new record is an ex-
pected range extension. We thank G. Davidse for the identification

St. George Parish: Morne Anglais trail at summit, S of Giraudel, 2 Jun 1996, Hill

28077 (atrc, ILLS, ES).

lsachne arundinacea (Sw.) Griseb. This is another clambering grass of exposed
wet mountain slopes, and its general range is Jamaica to Trinidad S to Bo-
livia and N to Mexico. In the Lesser Antilles it has been previously col-
lected on St. Kitts, Guadeloupe, and Grenada. We thank G. Davidse for
the identification.

Hitt AND JAMES, Plant records for Dominica S010

aul Parish: trail to summit of Morne Trois Pitons, N side, just NE of Pont Cassé,

9 = ie Hill 25340 (atrc, MO, VT).
Leptochloa virgata (L.) Beauv. A very widely distributed grass which ranges
from the southern United States through the Caribbean and Central America
to South America. It has been collected on many of the Lesser Antilles, in-
cluding nearby Marie Galante and Martinique, and was to be expected on
Dominica.

St. Peter Parish: Clement James’ property, Anse a Liane trail to coast, 0.9 km N of
Colihaut, 26 Feb 1997, Hill & Phillippe 28962 (atrc, ILLS, MO, TAES).
Pharus lappulaceus Aubl. This wide-ranging, wide-leaved grass is found in
forests from the Caribbean and Central America to southern South America.
Howard (1979) noted the similar P. ae Kunth from Dominica. We thank
T.S. Filgueiras for the identificatio

St. Mark Parish: Petit Coulibri, Morne Pie 15 Mar 1993, Hill 24796 (atrc, CLEMS).
Phragmites australis (Cav.) Trin. ex Steud. Probably due to the general lack
of suitable habitat, the common reed has not previously been collected on
Dominica. Otherwise, it is widespread in both tropical and temperate por-
tions of the world, and has been collected previously in the Lesser Antilles
on Guadeloupe, Martinique, and St. Lucia.

St. John Parish: marsh at N side of Coconut Beach Hotel, Prince Rupert Bay, Ports-
mouth, 29 May 1996, Hill 27993 (atrc, ILLS).
Spartina patens (Ait.) Muhl. This grass, usually found in salt marshes and
flats along the coast, was a surprising find in Dominica where these habi-
tats are essentially lacking. The species generally ranges from the E coast of
North America from Quebec to Mexico, Central America, and the Carib-
bean. In the Lesser Antilles it has been collected previously on St. Martin,
Antigua, and Guadeloupe. We thank D. Nicolson for the identification.

t. Andrew Parish: just N of the mouth of the Londonderry (Tweed) River, Londonderry

Estate, coastal rock ledges, 14 Dec 1993, Hill & James 25475A (atrc, VT).
Urochloa plantaginea (Link) Webster. Often reported as Brachiaria plantaginea
(Link) Hitchc., this grass of unstable areas ranges widely from Mexico and
the Caribbean to Brazil and Bolivia. As with Phragmites, it may be a recent
introduction, though Dominica falls within its general range. It has been
previously collected in the Lesser Antilles only on St. Lucia. We thank TS.
Filgueiras for the identification.

St. Paul Parish: Morne Daniel Road, Canefield, 10 Mar 1993, 24701 (atrc, CLEMS).

MAGNOLIOPSIDA

COMBRETACEAE
Conocarpus erecta L. The button mangrove (buttonwood) is widely distrib-
uted in the American tropics along the coasts of Florida, USA, S through

302 Sipa 18(1)

Mexico and Central America to the Caribbean and tropical South America
and it has also been collected in West Africa. Buttonwood has been found
on nearly all of the other Lesser Antilles. Hodge (1964) noted: “the com-
mon genera of mangrove (RAizophora, Avicennia, Laguncularia, and Conocarpus)
are absent from Dominica, a fact easily accounted for by the lack of suffi-
cient lowland sites on an island where the coastline is very precipitous.”
Diligent searching by Arlington James and other members of the Division
of Forestry and Wildlife have resulted in the verification of the occurrence
on Dominica of three of these genera, two of which (Avicennia and Conocarpus)
are reported here as new. Of the four genera listed by Hodge, only Rhizophora
has not yet been found on Dominica. Laguncularia was reported by Nicolson
(1991). Only one individual of Conocarpus has been found.

St. Andrew Parish: Ballyboo Beach, Batibou Bay, Hampstead, 14 Dec 1993, Hill &
James 25454. (atrc, CLEMS).
CONVOLVULACEAE
Convolvulus nodiflorus Desr. This species is widespread in dry scrub in the
West Indies, Central and South America. It has previously been collected
on the majority of the Lesser Antilles, including nearby Guadeloupe anc
Martinique, and was to be expected on Dominica. We thank D. Nicolson

—
tu

for the identification.
St. Joseph Parish: Morne Raquette eee Au Piton Road, 18 Dec 1993, Hi// 25526

(atrc, GH, MO, NY, US, USF). St. r Parish: S of Morne Espagnol between Bioche
and Colihaut, coastal highway, 7 Jun 1996, Hill 28248 (atrc, MO, US).
EUPHORBIACEAE

Euphorbia balbisii Boiss. A species endemic to the Lesser Antilles, also known
as Chamaesyce balbisii (Boiss.) Millsp., E. balbisii was previously known only
from Barbuda, Antigua, Guadeloupe, and La Désirade. It is a prostrate spe-
cles = wind-swept salt-sprayed coastal cliffs in Dominica.
ndrew Parish: Woodford Hill ie just E of Pre. La Soie E of an old quarry, 14

Be ee Hill & James 25464 (atrc, NY
FABACEAE
Dalbergia monetaria Lf. Generally distributed from Jamaica to tropical South
America, Howard (1988) reported that the species previously had been collected
in Guadeloupe, Martinique, and St. Vincent. Vélez (1957) reported this species
as present on Dominica, but Nicolson (1991) dismissed this as a misidentification.
Differing from D. ecastaphyllum (L.) Taubert (which is common on Dominica
and has unifoliolate leaves) by its pinnately compound leaves (3—S leaflets),
it is rather common locally along the banks of the Indian river. The identi-
fication was confirmed by V. Rudd (SFY).

St. John Parish: Indian River banks, ooo 3 Mar 1997, Hill & Phillippe 29056
(atrc, BRIT, ILLS, MO, NY, US, USF, VT

HILL AND JAMES, Plant records for Dominica 303

MALVACEAE
Sida jamaicensis L. This wide-ranging species is known from Mexico, the
Caribbean, Central America, and northern South America. In the Lesser Antilles
it has previously been collected on St. Barts, Antigua, Montserrat, Guadeloupe,
and Les Saintes in the dry scrub forests. We thank Paul Fryxell for the iden-
tification.

St. Joseph Parish: Morne Raquette (Rachette), Au Piton Road, 4 Jun 1994, Hi// 25808
(atrc, NY, TEX, USE).
NYMPHAEACEAE
Nymphaea amazonum C.Mart. & Zucc. This night-flowering water lily ranges
throughout the Caribbean and from Mexico to southern Brazil. It previ-
ously has been collected in Guadeloupe, Marie Galante, Martinique and St.
Lucia in the Lesser Antilles. Suitable habitat for the species is very scarce
on ae island. We thank J. Wiersema for the identification.

Andrew Parish: roadside 0.5 mi N of Melville Hall Airport, Londonderry Estate,

16 ane 1993, Hill 24844 (atrc, CLEMS).
OLEACEAE
Forestiera segregata (Jacq.) Krug & Urb. Another new discovery in the dry
scrub forests of western Dominica, this species ranges from the coastal SE
USA, S to the Lesser Antilles. It has been collected previously on Anguilla,

Barbuda, Antigua, La Désirade, and Marie Galante in the Lesser Antilles.
We thank A. Goldberg and D. Nicolson (US) for the determination.

St. Joseph Parish: Morne Raquette (Rachette), Au Piton Road, 18 Dec 1993, Hill 25550
(atrc, GH, NY, US).
RUBIACEAE
Randia nitida (H.B.K) DC. This spiny coastal shrub is distributed from Mexico
to South America and the Caribbean. In the Lesser Antilles it has been re-
ported from Martinique, St. Lucia, St. Vincent, and Grenada. It is distin-
guished from the more frequent R. aculeata L. by its flower size (3—3.8 cm
vs. 2cm long in R. aculeata) and fruit size (3.2—5 cm vs. 1-1.2 cm long in
R. aculeata). We thank D. Nicolson (US) for the identification.

St. Paul Parish: Rodney’s Rock on the coast, Hertford Estate, 13 Dec 1993, Hill 25425
(atrc).
RUTACEAE
Pilocarpus racemosus Vahl. This Caribbean endemic ranges from Cuba to the
Lesser Antilles. It has been collected previously on Montserrat, Guadeloupe,
and Meranicne in the Lesser Antilles and was to be expected on Dominica.
It is restricted to windswept coastal forests and is associated with the palm
Rhyticocos on the N coast of Dominica. We thank D. Nicolson for the iden-
tification.

304 Sipa 18(1)
St. Andrew Parish: cliffs at W side of Ballyboo Beach, Batibou Bay, Hampstead, 16
Mar 1993, Hill 24823 (atrc, CLEMS, NY, TAES, US, USF); same locality, 14 Dec 1993,
Hill & James 25453 (CLEMS).
VERBENACEAE
Avicennia germinans (L.) L. The black mangrove was a particularly interest-
ing find on Dominica. While it was thought to be absent by previous au-
thors (see comments under Conocarpus, above) it was located by the second
author in recent years at several places, the most unusual one in elevated
cliffside sedge-dominated seeps on the NE coast of the island. Other popu-
ations are restricted to a few square meters of level land at the mouth of
small rivers. Like Conocarpus, Avicennia is widely distributed along coasts
from Florida and Texas, USA, to northern South America. It has been found
on nearly all of the other Lesser Antilles.

St. David Parish: mouth of the Ouayaperi River, Corossol Beach, 0.6 km N of La Plaine,
12 Dec 1993, Hill 25390 (atrc, CLEMS, BRIT, ILLS, MO, NY, US, USF, VT). St. An-
drew Parish: Woodford Hill Estate, rast E of Pte. La Soie E of an old quarry in sedge
seeps, 14 Dec se Hill & James 25469 (atrc); mouth of Eden River, Eden Estate, S of
Crompton Point, 14 Dec 1993, Hill & James 25473 (atrc, GH, NY, US, USF).
VITACEAE
Cissus obovata M. Vahl. A vine endemic to the Caribbean, C. obovata is eas-
ily distinguished from the more common C. vertici/lata (L.) Nicolson & Jarvis
by its trifoliolate rather than simple leaves. It ranges from Cuba, S to the
northern Lesser Antilles. Collections previously have been made in the Lesser
Antilles on St. Martin, St. Barts, and Barbuda.

—

St. George Parish: head of trail to Trafalgar Falls, Trafalgar E of hydroelectric facility,

25 May 1996, Hill 27871 (atrc)
ACKNOWLEDGMENTS

We would like to acknowledge the continuing kind cooperation and support
of the Ministry of Agriculture of the Commonwealth of Dominica, and es-
pecially to the helpful staff of its Division of Forestry and Wildlife. Finan-
cial support for this research has been obtained primarily through various
units of Clemson University (particularly the Archbold Tropical Research
Center), Texas A&M University, and The University of Maine at Orono.
The senior author was also supported by grants from the Illinois Depart-
ment of Transportation (IDOT) and the Critical Trends Assessment Pro-
gram (CTAP). Special thanks are extended to the late John Archbold, whose
generosity made the research possible, to Dan Nicolson and his colleagues
at the Smithsonian Institution for their generous expenditure of time in
working with several of the identifications, to our other colleagues (cited
in the text) who assisted with identifications and collecting, and to Dr. Thomas
Lacher, former Director of the Archbold Tropical Research Center, for his
assistance and commitment throughout this study.

HILL AND JAMES, Plant records for Dominica 305
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A NEW ADDER’S-TONGUE (OPHIOGLOSSUM:
OPHIOGLOSSACEAE) FOR NORTH AMERICA

JAMES C. ZECH and PATRICIA R. MANNING
Department of Biology
Sul Ross State University
Alpine, TX 79832, U.S.A.

WARREN HERB WAGNER JR.

Department of Biolog)
University of Michigan
Ann Arbor, MI 48109, U.S.A.

ABSTRACT

The wides} sread Old World adder’s-tongue, Ophioglossum polyphyllu m A. Br. is reported
for the first time in North America. Formerly confused with RAG Pra nel, we now
recognize as O. polyphylfum collections from U.S.A. (Texas and Arizona) and
Oaxaca, San Luis Potosi, and Zacatecas). The earliest confirmed collection was in 1874 He

The two species are keyed out and illustrated, and specimens of O. polyphyllum are listed.
The known county distribution of the species in Texas is mapped, and the habitat and phenology
are briefly described.

RESUMEN
La lengua de serpiente del Viejo Mundo, muy ene OPA INE polyphyllum A. Br.
se Cita por primera vez en Norte América. P con O. engelmannii Prantl,

reconocemos ahora como 0. polyphyllum colectas de E.E.U.U. (Texas y Arizona) y Mexico
(Hidalgo, Oaxaca, San Luis Potosf, y Zacatecas). La primera colecta confirmada fue en 1874.
Se ofrece una clave de identificaci6on y se ilustran las dos especies, y se listan los especimenes

de O. polyphyllum. Se cartografia la distribucién conocida por condados en Texas, y se describen
brevemente el habitat y la fenologia.

The adder’s-tongues, Ophioglossum (Ophioglossaceae) are a small group
of striking and unusual species, with the highest known chromosome numbers
in the plant kingdom. The gametophytes are subterranean, the roots lack
root hairs, and the leaf is made up of two parts: the sterile blade portion
(trophophore) and the spore-bearing portion (sporophore). The trophophore
has complex reticulate venation and is believed to be phyllodial in origin
(Wagner 1979). The sporangia are deeply sunken in the apical portion of
the sporophore. There are probably as many as 35 species worldwide but
two areas, India and Africa, seem to be regions of maximum diversity. Many
of the species occur in open arid sites and remain underground until sea-
sonal or sporadic rains stimulate the leaves to push up through the soil and

Sipa 18(1): 307-313. 1998

308 Sipa 18(1)

grow to full size before discharging the spores. In America north of Mexico
seven onan species are traditionally recognized: Ophioglossum pusillam
Raf., O. vil ial ie . californicum Prantl, O. nudicaule L.f., O. petiolatum
= 0. rotalophoroides Walter, and O. engelmannii Prantl. However, we
have eney discovered another species heretofore unknown from the New
World. We originally interpreted che plants in question as a narrow form
of O. engelmannii (Zech & Manning 1996), but subsequent study reveals a
number of differences from that species, and we now identify it confidently
as O. polyphyllum A. Br. The earliest collection was in 1874. Clausen’s monograph
(1938) does not even recognize this species anywhere in the world; he merely
synonymized it with O. va/gatum. Nevertheless, we have come a long way
taxonomically from the first half of this century, not only in regard to
Ophioglossaceae, but Lycopodiaceae, Isoétaceae, Thelypteridaceae, and many
other pteridophyte groups. Lellinger’s (1985) manual of North America
pteridophytes does not list O. polyphyllum, nor does Flora North America
north of Mexico (Wagner & Wagner 1993

This species has an enormous range: Africa, Asia (e.g., S. China, India),
Polynesia (e.g. Hawaii), and now North America. It is probably made up of
a number of subspecies, but these are held together by a set of distinctive
characters, described below. There is no question that it is most closely re-
lated to O. enge/mannis. Its main point of resemblance involves the venation
pattern in which the major areoles include minor areoles, a condition com-
monly referred to as bireticulate. In most species the major areoles of adder’s-
tongues possess only free included veinlets that themselves do not inter-
connect. Large specimens of O. nuadicaule (the form known as O. e//ipticum
Hook. & Grev.) usually possess bireticulate veins (Wagner et al. 1984), as
does the Old World O. costatum A. Br. (Burrows 1996). Both Ophioglossum

vilgatum and O. reticulatum show a tendency toward bireticulation, but the
relatively few areoles are coarser and more open. The very fine bireticulation
in O. engelmannii and O. polyphyllum is quite distinctive.

Ophioglossum engelmannii is apparently a strictly New World taxon, and
limited to North America, including Mexico, and Central America. It is
commonly referred to as “Limestone adder’s-tongue’” for its tendency to grow
on soil layers above limestone. In the United States it usually appears dur-
ing and after rains, but it may reappear in the same places during later high
precipitation periods. Dean P. Whittier of Vanderbilt University has actu-
ally forced a new “brood” of leaves to form in late summer by watering a
site where the plants normally appear only in the spring (pers. comm.).

A key to the two species is presented below. To see the venation of a dried

on

specimen more clearly, a drop of 95% alcohol placed on the laminar surface
will, after soaking in, clear the leaf temporarily and will not damage the

ZECH, ET AL., A new Ophioglossum for North America 309

specimen. Collectors should be warned not to break off or remove the char-
acteristic elongate brown leaf sheaths that surround the green leaf bases:
collecting can be accomplished best by carefully digging up the specimens.
This has the added advantage of showing the roots, and their numbers and
relative sizes.
Roots 25(15—30), thinner, mostly 0.6(0.3—0.9) mm thick, yellow to dark brown,

commonly somewhat crooked (dried); persistent old leaf bases numero

(2-9); pag ener narrowly ovate, attenuate proximally and distally, ip

width 4, 4.8(3.2-—7.7) x 1.2(0.7-1. Ae cm; oe clusters 1.8(0.6—2.6)

x 0.25(0.2 —0.3) cm; apicula 1.0(0.3—1.8) m O. polyphyllum
Roots 15(6—24), thicker, 0.9-1.1(0.3-2. eA mm, we darker or blackish, usu-

ally straight (dried); persistent old leaf bases few 2(1—4); trophophore

broadly ovate to oblong, length/width = 3, 6.0(4.5-8.5) x 2.0(1.5—2. z: ae

gial clusters 2.5(2.0-4.0) x 0.25(0.2—0.4) cm; apicula 0.6(0.0

O. ae (Fig. 1)

The county distribution of Ophioglossum polyphyllum in Texas is shown in
Figure 2 (triangles) in comparison with O. engelmanni (dots). The map shows
a dramatic difference in range between the two species. Most of the collec-
tions of O. polyphyllum were made along U.S. Highway 90 or on roads nearby:
in Brewster County near Alpine and Marathon; in Jeff Davis County near
Valentine; and in Presidio County near Marfa. On the basis of our experi-
ence, we believe that an effort ought to be made to explore much further
afield in other Texas (as well as New Mexico and Arizona) counties, e.g.,
Hudspeth, Culberson, Reeves, Ward, Pecos, and Terrell in Texas, and also
areas to the south in Mexico (e.g., Coahuila, Chihuahua, and Sonora states
and southwardly adjacent states).

There are many reasons why O. polyphyl//um was not heretofore collected,
among the most important being its occurrence among grasses, the shape
of the leaf resembling some monocots, and especially the very short period
of appearance above ground, which after strong rains probably rarely ex-
ceeds two weeks, the latter week presenting only dying straw colored and
withered blades.

Plants are found in shallow ditches and troughs along the roadsides (Fig.
3). The adjacent landscape is open rangeland. While no plant species occur
consistently with O. polyphyllum, the two most prevalent are the grasses Buchloe
dactyloides and Bouteloua curtipendula. Other, less consistent associates are Bowteloua
gracilis, Chloris verticillata, Hilavia mutica, Panicum obtusatum, and Bothriochloa
sp. Among the forbs are Asclepias oenotheroides, Berlandiera lyrata, Cirsium
texanum, Cooperia drummondii, Croton Pottaii, Engelmannia pinnatifida, Ratibida
columnaris, and Thymophylla sp. The vegetation is typical Chihuahuan Desert
grassland, and includes such cactus genera as Coryphantha, Echinocereus, and
Opuntia. The substrate involves various clays that are usually deep and allu-

310 Sipa 18(1)

Fic. 1. A semi-diagrammatic comparison of O. polyphyllum (left) and O. enge/mannii (right).
ae bar = 2 cm

vial. Plants within adder’s-tongue colonies vary from several to hundreds.
Vegetative reproduction is accomplished by horizontal roots, the prolifera-
tions producing plantlets as far as ten centimeters from the genet.

Our records outside of Texas are very few, and all were previously identi-
fied as O. engelmanni?. In Arizona it was found in “mesas around the Mus-
tang Mts.” The habitats in Mexico (what little is recorded on the labels)
appear to be more or less similar to those in Texas. The Rollins and Tryon
collection listed below was found in “clay soil over limestone, in shade of
small arborescent Opuntia in Larrea and Prosopis desert,” and the Reeder et

——

ZECH, ET AL., A new Ophioglossum for North America a

esse,
| “St
a a woe :

Fic. 2. Presently confirmed county records in Texas for O. polyphyllum (triangles) and O.
engelmannit (dots).

al. collection in “thorn savannah in the open among Eachloé dactyloides and
Bouteloua stolonifera.” The Pringle and Purpus Mexican collections lack habitat
data.

ollection citations: U.S.A. ARIZONA: Mustang Mountains, 13 Sep, C. G. Pringle in
1884, (GRAY). TEXAS. (Collections made along Hwy 90 and ee in SRSC, unles
otherwise indicated. Brewster Co.: 5.5 mi E of Marathon N of Hwy 90, 7 Jun 1995, Newie
896; 5.9 mi W of Marathon S of Hwy 90, 20 Jun 1995, et oe 6.5 mi E of Alpine
S of Hwy 90, 8 Jul 1995, epee 914; 12 mi W of Marathon S$ of Hwy 90, 5 Jul 1995,
Manning s.n.; 6.5 mi W of Marathon N of Hwy 90, 24 Jun 1996, ee 959;5.7 mi E
of Marathon, 11 Jun 1995, A.M. Powell G S.A. Powell 6085; 6.0 mi E of Marathon N of
Hwy 90, 13 Jun 1997, Manning 1013; 17 Jun 1997, Manning & Zech Bee (MICH); 24
Jun 1997, Manning & Zech 1017B (MICH); 6.2 mi E of Marathon S of Hwy 90, 14 Jul
1997, Hansen et al. 40; 6.3 mi S of Brewster/Pecos Co. line, 30 Jun 1997, Pee 1034.
Jeff Davis Co.: 7.8 mi W of Valentine E of Hwy 90, 11 Jul 1995, Manning 915; 25 Jun
1997, Manning & Zech 1020 (MICH); 0.2 mi SE of 505/166 jct. on loop 166, 14 Aug 1971,
Keil 1699 (GRAY); 25 Jun 1997, Manning & Zech 1031 (MICH). Presidio Co.: 0.1 mi E
of Marfa, 26 Jul 1995, Manning 925.

Sipa 18(1)

We
—
bo

Fic. 3. Habitat of 0. polyphyllum in shallow roadside ditch. TX: Jeff Davis Co., along loop
166

MEXICO. Hidalgo: Hill, near El Gallo, 7000 ft, 17 Aug 1902 C.G. Pringle (GH). Oaxaca:
Los Maranjos, vicinity of San Luis Tultitlanapa, Puebla, near Oaxaca, May 1908, C.A. re
3152 (GH). San Luis Potosi: 15 mi NE of San Luis Potosi, 6500 ft, 4 Sep 1961, /.R.
Reeder, C.G. hee & TR. Soderstrom 3300 (GH). Zacatecas: 32 mi N of Fresnillo, 25 Nov

1958, 7000 ft, R.C. Rollins and R.M. Tryon (GH).

There is no easily projected certainty about when the plants will appear.
Continuous drought will prevent the leaves from expanding and extending
above the soil. However, the best months in general are June and July, whenever
several rains occur. Apparently, chance rainy periods at any other time, however,
will bring forth an “out-of-season” growth, and we now have records from
May, August, September, October, November, and December. Our impression
is that O. polyphyllum “lies in wait” for rain, emerges through tightly com-
pacted soil that has been softened by the water, and then has a fairly short
time span to reproduce and disperse spores. However, based upon green-
house observations, even when water availability is not a factor, trophophores
will tend to die back following spore production and dispersal.

It should be noted that O. polyphyllum is ecologically a “loner,” in the
sense that it does not occur microsympatrically along with other species of
Ophioglossum (or Botrychium). Other Ophioglossaceae are noted for forming
“genus communities,” members of the same or closely related genera that

ZECH, ET AL., A new Ophioglossum for North America a135

co-occur syntopically (Wagner & Wagner 1983). To illustrate the usual situation,
in the United States in the region from the Carolinas to as ane Sonrhward,
we find O. nudicaule, O. crotalophorvides, and O. petiolatum (p eous
Botrychium biternatum and B. lunarioides) bog ties in ne same habitat, s sometimes
side by side and intermixed with each other. But this is evidently not the
case with O. polyphyllum according to Burrows (1996) who writes that in
Africa O. polyphyllum is “The one species of Ophiog/ossum that is rarely found

as a component of genus communities.”

ACKNOWLEDGMENTS

This study could not have been completed without the help of several
herbaria, including Gray Herbarium, Harvard (GH), Lundell Herbarium
(LL), University of Michigan (MICH), University of California Berkeley (UC),
New Mexico State (NMS), Sul Ross State (SRSC), Southwest Texas State
(SWT), Texas Tech (TTC), University of Texas at El Paso (UTEP), Botani-
cal Research Institute of Texas (BRIT), Texas A&M University (TAMU),
and the help of Christopher Ruggia who kindly provided the distribution
map. Alan R. Smith and R.D. Thomas made useful suggestions for the
manuscript.

REFERENCES

Burrows, J.E. 1996. The genus Ophioglossum L. in southern central Africa. In: J.M. Camus,
M. Gipsy, and R.J. Jouns, eds. Pecticologict Perspectives. Royal Bot Gard. Kew. Pp.329-336.

CLAUSEN, RT. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club. 19:1—177
Ciute, W. 1901. Our ferns in their haunts. F. ee Co. NY.
LELLINGER, D.B. 1985. A field manual of the ferns & fern allies of the United States & Canada.

Smithsonian Institution Press, Washington, DC.

WAGNER, W.H. 1979. Reticulate veins in the systematics of modern ferns. Taxon 28:87—95.

Wacner, W.H., C.M. ALLEN, and G.P. Lanpry. se i eee Hook. & Grev.
in Louisiana ad the taxonomy of O. nudicaule L. f. Castanea 49:99—

WAGNER, W.H. and ES. WacGNner. 1983. Genus communities as a systematic tool in the
study of New World Botrychinm (Ophioglossaceae). Taxon 32:51—-63.

WAGNER, W.H. and E.'S. Wacner. 1993. Ophioglossaceae C. Agardh. Adder’s-tongue Family.
In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America
north of Mexico, 3+ vols. New York and Oxford. Vol. 2, pp. 85-106.

ZECH, J.C. and P.R. MANNING. 1996. Ophioglossum engelmannii in Trans-Pecos, Texas. Amer.
Fern J. 86:102.

a

NOTES ON THE FLORA OF TEXAS WITH ADDI-
TIONS AND OTHER SIGNIFICANT RECORDS
LARRY E. BROWN
Houston Community College

1300 Holman, Houston, TX 77007, U.S.A.

and
Spring Branch Science Center Herbarium
88506 Westview Drive
Houston, TX 77055, U.S.A. plantman@ flash. net
STUART J. MARCUS
Trinity River National Wildlife Refuge

Liberty, TX 77575, U.S.A. r2rw_tr@fws. gov

ABSTRACT

The following 15 taxa are documented new to Texas: A/rer lec sessilis, Cyperus eragrostis
Urochloa ramosa, Lindernia crustacea, Phyllanthus fraternus, Stellaria parva, Alysicarpus vaginalis,

Cardamine debilis, Rhynchospora debilis, Lilaeopsis chinensis, Rime paraguayensis, intl
coarctatum, ee es Ls meisnertanum, and Cuscitta pe. Sls a Significa
new collections records are provided for Scirpus cubense, Polyg ‘ Phyllentbus
nirurt, Ludwigia soared ees ins Data are presented to aiecnn the presence
of Eleocharts compressa in Texas. Taxa are listed oe y by family, genus, and species
with annotations and citation of voucher specimens

RESUMEN

A ]y. 7) “aerssse Cy 4 - rae

Los siguientes 15 taxa se citan como nuevos para Tex:
Urochloa ramosa, Lindernia crustacea, Phyllan ‘bus raters, Stellaria parva, ap eae vaginalis,
Cardamine debilis, Rhynchospora debilis, Lilaeopsis chinensis, Rumex xX paraguayensts, Saccharum
coarctatum, uF ee eas. Polygonum meisnerianum, y Cuscuta polygonorum. Se ofrecen nuevas
s importantes para cinco: Scirpis cubense, Polygontum cespitosum, Oca niruri, Ludwigia
microcarpa, y Armoracia ees Se presentan datos para cuestionar la pr a de Eleocharis
compressa en Texas. Los taxa estan listados alfabéticamente por famnili ia, género, y especie

~

con anotaciones y ie de los pliegos testigo.
INTRODUCTION

Floristic work in the Houston area has resulted in a mimeographed plant
checklist for: Brazos Bend State Park, Fort Bend County by Larry E. Brown,
Frank Gregg, and Kay Lewis; Davis Hill State Natural Area, Liberty County
by Larry E. Brown, Charles D. Peterson, and Joe Liggio; and the entire Houston
area (Harris and adjacent counties) by Larry E. Brown. In addition, the present
authors have just completed a plant checklist for the Trinity River National

Sipa 18(1): 315-324. 1998

316 Sipa 18(1)

Wildlife Refuge, Liberry County. This work coupled with plants sent to
the SBSC herbarium for identification has revealed a number of plants new
to Texas and significant new records for others. Some of these records were
provided to Stanley D. Jones prior to the publication of his book, Vascular
Plants of Texas (Jones et al. 1997). Here we give data to support inclusion of
these plants in this new publication.

Unless otherwise indicated, all collections cited in this paper are in the
Spring Branch Science Center Herbarium (SBSC), Houston, Texas. Dupli-
cates of some collections are at TEX or TAES. Herbarium acronyms follow
Holmgren et al.(1990).

AMARANTHACEAE

Alternanthera sessilis (L.) DC.—A few plants of this pantropical and federal
noxious weed were found growing in the bottomland of the Trinity River
in the Davis Hill State Natural Area; 4 Jul 1994, Brown 18040. Clewell
(1985) reported it for Escambia County in the Florida Panhandle and Tho-
mas and Allen (1996) mapped it for ten, mostly central, Louisiana parishes.
Nelson et al.(1997) reported it new for South Carolina and cited published
reports for its occurrence in Alabama and Mississippi. The key in Mears
(1977) can distinguish the Texas species of A/ternanthera, including A. sessi/7s.

APIACEAE

Lilaeopsis chinensis (L.) Kuntze—Affolter (1985) mapped this species along
the Atlantic and Gulf Coast from Nova Scotia south to Florida and west-
ward along the Gulf Coast to Louisiana (one collection). Thomas and Allen
(1996) mapped it for eight coastal parishes west to Vermillion Parish. The
only Texas collection is from Galveston Bay below the Abshire House in
the Abshire Wildlife Management Area at Smith Point, Chambers County,
7 May 1993, Brown 16926. These plants were submerged at high tide and
exposed during low tide.

BRASSICACEAE

Armoracia lacustris (A. Gray) Al-Shehbaz & Bates (lake cress)—Correll and
Johnston (1970) did not report this white flowered aquatic crucifer for Texas.
The first record appears to be in Al-Shehbaz and Bates (1987) where it is
mapped in Tyler County. We found it to be infrequent in standing water in
the Davis Hill State Natural Area, Liberty County (31 May 1996, Brown &
Ligeto 19172) and in the bottomland of the Trinity River National Wild-
life Refuge, Liberty County (26 Apr 1997, Brown & Marcus 20262). These
plants produced numerous flowers but had poor fruit development. Mo-
lecular studies based upon DNA (Les 1994) indicate that lake cress should
be placed into a monotypic genus Neobeckia as N. aquatica (Eaton) Greene
and that the cause for a poor seed set is suggested by the discovery of a trip-
loid chromosome number of 27 = 28 (Les et al. 1995).

Brown AND Marcus, Notes on the flora of Texas 317

Cardamine debilis D. Don—Plants tentatively identified as this non-native
species have been collected in and around nurseries, greenhouses, and yards in
the Houston area. These are similar to the native C. pennsylvanica Willd. and
another introduced species mostly present in the norteast U.S. and Canada, C.
flexuosa Withering. Rollins (1993) separated C. debilis from the other two species
by its fibrillose roots and fruits <1 mm wide. Botanists may wish to reex-
amine their collections, especially those identified as C. pennsylvanica, to see if
they may fit the alae of C. debilis. It also would be helpful to search for
additional charac te these three similar species. Lipscomb (1978) cited
the Shinners collection as a Ceoahe: for the presence of C. pennsylvanica in Texas.
Voucher specimens: Dallas Co.: weed under lath, mostly along walks in clay, Northaven
Gardens, north Dallas, 6 Nov 1960, Shinners 29154 (TEX). Dewitt Co.: in Sandies Creek,
2 mi iB of Westhoff, 5 Mar 1995, D. Muschalek s.n. (SBSC). Galveston Co.: weed in nurs
n Kemah, | Apr 1975, Waller & Bauml 3460 (TEX). Harris Co.: ie around ie
pe in the Mercer Arboretum and Botanical Bee off of Aldine Westfield Rd,
1 Nov 1997, Brown 21715 (SBSC); growing in pots at the Edith Moore Sanctuary off of
Memorial Avenue in Houston, 27 Mar 1992, Brown ae (SBSC); weed at edge of San
Jacinto Bldg at ae Central College, 1300 Holman Street in Houston, 25 Nov 1997,
Brown 21742, (SBS
CARYOPHYLLACEAE
Stellaria parva Pedersen—Landry et al. (1988) reported this chickweed new
to North America from Acadia and Jeff Davis Parishes in Louisiana. Landry
et al.(1989) provided a more complete discussion of its presence in Louisi-
ana and cited an additional collection from St. Landry Parish. Thomas and
Allen (1996) added Lafayette Parish to the Louisiana distribution. We dis-
covered this species on a few disturbed sandy sites in the Trinity River National
Wildlife Refuge, Liberty County, 11 Mar 1997, Brown & Marcus 20104; 6
Apr 1997, Brown & Marcus 202106. Since the original description is from
Argentina in 1961 and the first Louisiana collection is in 1966, Landry et
al. (1989) suggest it may have arrived in Louisiana and Argentina from an
unknown native source. However, they failed to note that Pedersen (1961)
in the protologue reported collections from Argentina and Paraguay in 1869
and 1872 and up to 1957
CUSCUTACEAE (CONVOLVULACEAE)
Cuscuta polygonorum Engelmann—Correll and Johnston (1970), Hatch et
al.(1990), Johnston (1990), and Jones et al. (1997) all include this dodder
for Texas, apparently from Yuncker (in Lundell 1943), who included it fide
mall. Even though Small (1933) gave its range from Maryland to Texas,

the first Louisiana collection of it was in 1979 with a second collection in
1981 (Gandhi & Thomas 1983; Gandhi et al. 1987). The first Texas collec-
tion appears to be from Brazos County where it was growing on smartweed
in the first bottomland along Highway 30, east of the Highway 6 bypass of
Bryan/College Station, Sep 1996, Cheatham, Marshall & Jones s.n.

318 Sipa 18(1)

CYPERACEAE
Cyperus eragrostis Lam.—Tucker (1987, 1994) reported that this species has
become naturalized in southeastern Texas but he cited no collections nor
furnished details about its status in Texas. Tucker (1987) indicated it is native
to the Pacific Coast of the U.S., a waif in South Carolina, and naturalized in
southern Europe. It is similar to C. ochracews Vahl and may be under this
name in other Texas herbaria. The key in Denton (1978) can separate these
two taxa and other similar species. The larger number of collections from
LaPorte near the Barbours Cut Terminal of the Port of Houston suggest it
may have arrived in hes — area by ship.

oucher ace Galveston Co.: ca. 100 m from Gulf of Mexico on East Beach in
Galveston, | Jul | 981, pe (TAES). Harris Co.: in roadside ditch of echo
Rd, ca. 0. mi S of Hwy 225,18 Jun 1983, Brown 7207 (SBSC); in the Marshall Tra
the Co ompaq Computer Corp at the intersection of Hwy 249 and Louetta Rd, S of ce
23 May 1992, Brown 16104 (SBSC); weedy plants in the Park Harbor development along
Barker-Cypress Rd N of Addicks Reservoir, W of Houston, May 1994, collector unknown
(SBSC); a "single ylant in a prairie at end of Ramsey Rd, S of Keith Rd, E of Beltway 8 in
Pasadena, | Jun 1994, Brown 17952 (SBSC); small population on verge of Barbours Cut
Rd between hwy 146 and Barbours Cut Terminal on Galveston Bay in LaPorte, 16 Jun
1994, Brown 18022 (SBSC); frequent in ditch along Hwy 146 on W side across from en-
trance to Barbours Cut Rd in LaPorte, 27 Jun 1994, Brown 18023 (SBSC); ee plant at
the Texas Parks and Wildlife tease office, near business Hwy 146 in LaPorte, 24
Jan 1995, Brown 18403 (SBSC); in low waste areas along Southern Pacific ae Trac ts
just E of Hwy 146 in LaPorte, S of the Baytown Tunnel and Bridge complex, 19 July 1995,
Brown eae (SBSC).
Eleocharis compressa Sullivan—Correll and Johnston (1970) separate E.. compressa
and EF. acutisquamata Buckley. by the wider compressed culms in E. compressa
and the more narrow non-compressed culms in E. acutisguamata. Further, they
reported FE. compressa only for San Augustine County in East Texas and re-
stricted E.. acutisquamata to the Edwards Plateau, the north cental, and southeast
(Refugio County) portions of the state. In addition, they indicated (page 275)
that they are probably conspecific which is suggested by the overlapping culms
width measurements used to separate them in their key. We were unable to
borrow Texas and non-Texas material of both entities from TEX for they are
out on loan to $. Galen Smith for the Flora of North America project. How-
ever, in a survey of collections at SBSC and ASTC, we found only specimens
referable to EL. acutisquamata Buckley in East Texas (San Augustine and Sabine
counties). From the specimens available to us, both entities can be readily
separated by the nearly round culms (7—9 angled) of E. acutisquamata that are
up to 25 cm tall and 0.5 mm wide in contrast to the flat culms of E. compressa
that are up to 42 cm tall and 2 mm wide.

Rhynchospora debilis Gale—Gale (1944) described this taxon as a new spe-
cies and noted its similarity to R. fascrcularis (Michx.) Vahl. Both of these

BROWN AND Marcus, Notes on the flora of Texas 319

species were growing together at the edge of Galveston Bay on a sandy bench
below the bluff next to the Abshire House in the Abshire Wildlife Man-
agement Area at Smith Point, Chambers County, 7 May 1993, Brown 16932;
27 May 1996, Brown G Liggio 19138. Here they may be distinguished by
the taller (= 35 cm), thicker culms (= 1.5 mm wide at culm base), and flat
leaves (= 1.0 mm wide) of R. fascicularis in contrast to the shorter (ca. 15
cm) filiform culms (= 1 mm wide at culm base) and filiform leaves (= 0.5

wide) of R. debilis.

Scirpus cubensis Poeppig & Kunth [Oxycaryum cubense (Poeppig & Kunth) Lye}—
The first Texas collection of this sedge was in 1958 from Eagle Nest Lake
in Brazoria County, Hotchkiss 7653 (TEX). The second Texas collection is
from Fort Bend County where is was somewhat frequent in a floating mat
of vegetation in Elm Lake at Brazos Bend State Park, 7 Nov 1997, Brown
21737. Thomas and Allen (1993) mapped it for eight central and eastern
Louisiana parishes, none near the Texas border.

EUPHORBIACEAE

Phyllanthus fraternus G.L. Webster—The first Texas collection is from downtown
Houston growing on bare ground among planted shrubs; 17 Sep 1995, Brown
18753. Webster (1970) indicated it is native to Pakistan and India and spo-
radically introduced into Africa and America. He cited only three U.S. col-
lections, one from an abandoned field in Seminole County, Florida, and the
other two from New Orleans, Louisiana (including the first U.S. collection).
It is now mapped for nine Louisiana parishes mostly in the New Orleans area
(Thomas & Allen 1996). It may also expand its range in the Houston area.

Phyllanthus niruri L. subsp. lathyroides (Kunth) G.L. Webster—Correll and
Johnston (1970) indicated this taxon may be extinct in Texas (the only U.S.
records for this widespread tropical American species are from Texas) for
they reported it only from the Ottine Swamp in Gonzales County, prob-
ably based upon the 1935 Tharp collection at TEX. The only collections
since 1935 are the following.

Dewitt Co.: a group of plants ca. 2 ft tall, on sand under bridge on Thomaston Rd, W
of Hwy 87 along the Guadal He River, ca. 2 mi W of Thomaston Community, 11

1994, Muschalek s.n.(SBSC). Fayette Co.: S bank of the Colorado River at foot of bluff, .
of Monument Hill State ii Site, 11 Oct 1987, Carr 7867 (TEX).

The only known U.S. collections are two in the nineteenth century by
Wright, one of which is from the Colorado River bottomland, the one in
1935 by Tharp, and the two recent ones reported here.

FABACEAE
Alysicarpus vaginalis (L.) DC.—Isely (1990) mentions this native from the
Old World tropics as occurring in Texas. He has examined Fabaceae collec-

320 Sipa 18(1)

tions at TEX and probably discovered this Hardin County specimen, 4 mi
S of Kountze along roadside, 22 Sep 1960, McLeod s.n. We have collected
plants of this legume along a pipeline right-of-way in the Trinity River
National Wildlife Refuge, Liberty County, 13 Oct 1996, Brown & Marcus
19731.

LENTIBULARIACEAE

Utricularia foliosa L.—This large flowered, robust, floating bladderwort was
not reported for Texas by Correll and Johnston (1970), possibly because Small
(1933) indicated that it ranged from Florida to Louisiana. The first report
seems to be in Taylor (1989), where it is listed for North Carolina, Georgia,
Florida, Mississippi, Louisiana, and Texas. However, Taylor does not indi-
cate its distribution in Texas nor provide specimen citations. The majority
of the TEX and LL specimens were identified as the robust U. vulgaris L.
Taylor (1989) considers the North American representatives of U. valgaris
a distinct species, U. macrorhiza Le Conte, and indicates its absence from
eastern Texas and most of the southeastern U.S. Coastal Plain.

Voucher specimens: Brazoria Co.: floating aquatic in water in the Brazoria National
Wildlife Refuge S of Angleton, 26 Apr 1992, Brown 16020 (SBSC); Muldoon Club be-
tween Austin Bayou and Danbury, 19 May 1971, Fleetwood 10029 (TEX). Chambers Co.:
present in Anahuac National Wildlife, Jul 1990, Neaville s.n(SBSC). Fort Bend Co.: Brazos
Bend State Park in 40 Acre Lake beside dike between 40 Acre Lake and Pilant Lake, M

1994, Gregg s.n(SBSC); 29 May 1993, Gregg s.n. (SBSC). Hardin Co.: 28 Jun 1952, Tharp
& Tyson s.n(TEX); Ariola Rd, S of Kountze, home of Jack Lloyd, 9 May 1957, Whitehouse
32581 (TEX). Henderson Co.: Code Lake, 27 May 1940, mai 2000 (TEX). Jefferson
Co.: floating in canal along rd,4 mi N of Port Authur, 19 Jul 1946, Correll 13357 (LL);
J.D. seas Wildlife Mangement Area, 1-10 mi SW of Pore Arthur, Big Hill Bayou,
submerged, 20 Oct 1966, West 127 (TEX). Liberty Co.: floating on water in bayou about
7 mt E of Cleveland, Rte. 321, 12 Sep 1968, Correll 36458 (LL).

ONAGRACEAE

Ludwigta microcarpa Michx.—Bridges and Orzell (1989) reported this spe-
cies new to Texas from two roadside collections in Hardin County. They
state these collections are within former extensive wetland pine savannahs
on the Montgomery and Beaumont Formations of the Pleistocene Epoch.
A new county collection, perhaps from a younger formation, is from Chambers
County where it was frequent along the sandy roadside of Highway 562 ca.
2.8 mi east of the road to Robbins Point Park, 27 May 1996, Brown & Liggio
LOf52.

The following collections at SBSC complete the documentation of this
species in Texas. These roadside collections suggest it may be spreading westward

via highways.
Voucher specimens: Hardin Co.: frequent in a dry roadside ditch along hwy 69/287 ca.
mi N of Kountze, associated with Rhynchospora divergens and Fuirena breviseta, 2 Aug 1986,

Brown 10581, frequent in a roadside ditch at edge of a sandy acid area along Hwy 770, ca.

BROWN AND Marcus, Notes on the flora of Texas 321

0.4 mi W of intersection with Hwy 1003 between Saratoga and Kountze, 16 Aug 1986,
Brown 10641; on acid sandy soil with many bog plants along Hwy 421 ca. 3 mi E of inter
section with Hwy 326 N of Sour Lake, 9 Jun 1996, Brown oe
POACEAE
Saccharum coarctatum (Fern.) R. Webster—Webster and Shaw (1995) did not
consider the presence (Erzanthus) or absence (Saccharum) of an upper lemma
awn sufficient to separate these two genera. They, therefore, merged Evianthus
under Saccharum and made the appropriate combinations under Saccharum.
They did not cite any Texas collections of 8. coarctatum, possibly because
they did not examine sufficient material from Texas herbaria (the only Texas
herbarium cited is BRIT). The following collections at SBSC document its
presence in the state. The senior author first identifed these plants as E.
brevibarbis Michx. and then annotated them to S. brevibarbis (Michx.) Pers.
var. brevibarbis. Joseph K. Wipff, then at TAES, identified them to S. coaretatiin.
esas specimens: Liberty Co.: tall plants in moist ditch along Hwy 90, 5.4 mi E of
y Ol in Devers, 25 Sep 1993, Brown 17493; tall plants with brownish Me
ad Hwy 90, 4.9 mi E of Hwy 61 in Devers, 7 Nov 1994, Brown 17618. Tyler Co.: large
colony on low moist acid soil within Kirby State Forest off of Hwy 69, S of Warren, 8 Oct
1983, Brown 6674.

Uvochloa ramosa (L.) Webster [Panicum ramosum L.; Bachiaria ramosa (L.) Stapf}—
This native of tropical Asia has been planted in Texas and other states for
erosion control and wildlife food but has not been included in any Texas
floristic publications. It is reported for 31 Louisiana Parishes (Allen 1992).
An adventive Texas collection is from Chambers County where it was growing
at the edge of the Highway 1663 bridge over Whites Bayou northeast of
Hankamer, 25 Sep 1993, Brown 17488
POLYGONACEAE
Polygonum cespitosum Blume—Carr and Hernandez (1993) reported this Asian
species new to Texas (first collection from Jasper County in 1992) using
the spelling caespitosum. We are following the spelling in Kartesz (1994).
We discovered this species on moist to dry disturbed sites in the Trinity
River National Wildlife Refuge. Here the reddish flowers stand out in contrast
to the whitish flowers of P. Aydropiperoides Michx. and other similar species.
Voucher specimens: Liberty Co.: 15 Sep 1996, Brown 19701; 13 Oct 1996, Brown &
Marcus 19732; 9 Nov 1996, Brown 19787; 21 May 1997, Brown 20445. The following
two collections, ae! identified as P. persicaria L., are at TAES: Galveston Co.: Galveston
Island State Park, 15 May 1997, Rosen 0316. Tiber Co.: ca. 4.5 mi S of Dayton along
Hwy 146, 21 Oct 1987. oe 16050.

Additional state collections may be masquerading under this name in
other Texas herbaria.

sal meisnerianum Cham. & Schlecht. var. beyrichianum (Cham. & Schlecht.)

sn.—Correll and Johnston (1970) indicated this species has been re-

322 Sipa 18(1)

ported for Texas but saw no specimens. Park (1988) also saw no Texas col-
lections but reported U.S. collections from Florida and South Carolina and
from Plaquemines and St. Tammany Parishes in Louisiana. The only Texas
collection known to us is from Orange County where it was growing in a
slight swale under a canopy of Chinese tallow trees adjacent to a fallow rice
field near Nederland, Know/es s.n. The senior author originally identified
this specimen as P. sagittatum L. and additional collections may be filed under
this name in Texas herbaria.

Rumex paraguayensts Parodi—This native of southern South America (natu-
ralized in Florida and Louisiana) is now in Texas (Chambers County, present
in the East Unit of the Anahuac National Wildlife Refuge off of Hwy 1985,
SE of Anahuac, Spring 1991, Nev7/le 274). The key in Godfrey and Wooten
(1981) can separate this species from R. obovatus Danser, a similar natural-
ized South American species in northern Florida and Louisiana. Thieret (1969)
reported both species new to North America and speculated that R. obovatis
almost certainly occurs in Texas because one collection of it was made within
150 feet of the Texas border. However, based on Godfrey and Wooten’s key,
our specimen is R. paraguayensis.
SCROPHULARIACEAE
Lindernia crustacea (L.) KF. Muell.—This Indo-Malaya native ranges in the
U.S. from South Carolina, south to Florida, and along the Gulf Coastal Plain
to Louisiana (Godfrey & Wooten 1981). Vincent (1982) mapped it for 15,
mostly south central and eastern, Louisiana Parishes. He indicated it is spreading
westward and that it may eventually appear in East Texas. We found this
short blue flowered species on two disturbed sites in the Trinity River National
Wildlife Refuge, Librty County, 29 Sep 1996, Brown & Marcus 19703; 13
July 1997, Brown 20581.
ACKNOWLEDGMENTS

We wish to thank the following: Tom Wendt (TEX), James VanKley (ASTC)
and Stephan Hatch (TAES) for permitting access to their collections, for
the loan of herbarium material, and for sending copies of label data from
others; Guy Nesom and an anonymous reviewer for their careful reading
and improvement of the paper; all of the Houston area and state-wide ama-
teur botanists who have added collections to our local herbarium (SBSC)
and for their help in numerous other ways; and the Texas Parks and Wild-
life Department and local National Wildlife Refuges for permitting col-
lections of plants from their respective lands. Work on the Trinity River
National Wildlife Refuge was supported by Challenge Cost-Share Agree-
ment with the refuge.

BROWN AND Marcus, Notes on the flora of Texas 323

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=

. Thesis. The University of South-

CYPERUS FUSCUS (CYPERACEAE), NEW TO MIS-
SOURI AND NEVADA, WITH COMMENTS ON ITS
OCCURRENCE IN NORTH AMERICA

PAUL M. MCKENZIE BRAD JACOBS
U.S. Fish and Wildlife Service Missouri Department of Conservation
608 E. Cherry St. Room 200 P.O. Box 180

Columbia, MO 65201, U.S.A. —_ Jefferson City, MO 65102-0180, U.S.A.

CHARLES T. BRYSON
USDA, ARS, Southern Weed Science Research Unit
P.O. Box 350
Stoneville, MS 38776-0350, U.S.A.

GORDON C. TUCKER RICHARD CARTER
Stover-Ebinger Herbarium Herbarium, Department of Biology
Botany Department, Eastern Illinois University Valdosta State University
Charleston, IL 61920-3099, U.S.A. Valdosta, GA 31698, U.S.A.
ABSTRACT

Field and herbarium studies have documented Cyperus fuscus as new for Missouri and
Nevada. Localities, habitat data, lists of associated species, description, illustrations, photo
of habit, and discussion of weedy potential are presented.

RESUMEN

Los estudios de campo y de herbario han documentado Cyperus fuscus como nuevo para
Missouri y Nevada. Se aportan localidades, datos del habitat, listas de especies asociadas,
descripcién, ilustraciones, foto del habito, y discusién de su potencial como mala hierba.

INTRODUCTION

Cyperus fuscus L. [section Fusei (Kunth) Clarke} is native to Eurasia and
the Mediterranean Region of northern Africa, extending from Greenland
and Iceland to China south to Spain, Iran, Egypt, Algeria, and northern
India (Kiikenthal 1936; Tutin et al. 1980). The common name for this spe-
cies has been listed as “galingale,” “brown galingale,” or “black galingale”
(Weedon & Stephens 1969). It apparently was first discovered in North America
based on specimens collected by Herbert A. Young along Revere Beach in
Essex County, Massachusetts in 1877 (Knowlton et al. 1911; McGivney
1938). Cyperus fuscus was subsequently documented in California (McGivney

Sipa 18(1); 325-333. 1998

326 Sipa 18(1)

1938; Tucker 1993), Connecticut (McGivney 1938; Dowhan 1979; Tucker
1995), Maryland, (Kiikenthal 1936), Nebraska (Weedon & Stephens 1969;
Kolstad 1986; Rolfsmeier 1995), New Jersey (Britton 1886; McGivney 1938),
Pennsylvania (Rhoads & Klein 1993), South Dakota (Weedon & Stephens
1969), and Virginia (Hitchcock & Standley 1919; Kolstad 1986; Tucker
1987). Although the species has also been recorded for New York (e.g., Weedon
& Stephens 1969; Kolstad 1986), no specimens have been located and the
only ones so identified were actually C. diandrus Torr. (Mitchell & Tucker
1997). Although C. fuscus has been present on the North American Conti-
nent since at least 1877, its spread apparently has been slow, and outside of
brief accounts related to its increase in distribution, there has been little
attention given to it in the New World literature.

Given the great attention to new records of Cyperus in North America by
Charles Bryson, Richard Carter, Stanley Jones, Barney Lipscomb, Gordon
Tucker, and others in the last 20 years (e.g., Lipscomb 1978, 1980; Tyndall
1983; Carter et al. 1987; Carr 1988; Carter 1988; Sundell & Thomas 1988;
Carter & Bryson 1991; Webb et al. 1991; Bryson & Carter 1992; Jones et
al. 1993; Bryson & Carter 1994; Tucker 1994; Bryson et al. 1996; Carter et
al. 1996), it is interesting and somewhat amazing that the species has not
been discovered at more localities throughout the United States, especially
as Weedon and Stephens (1969) noted that it was a weed of rice fields in
the Old World. Outside the United States, the species has been discovered
in Canada (Gillett 1970), but it has not been found in Mexico (Tucker 1994)
or Central America (Tucker 1983).

—

DESCRIPTION

The following is a description of C. fuscus based on Kiikenthal (1936),
McGivney (1938), Fernald (1950), Kolstad (1986), and our observations.
Cyperus fuscus (Figs. 1, 2): a caespitose annual with few to numerous culms
and reddish-brown fibrous roots; culms upright, inclined, or decumbent,
2—30 (rarely 50) cm long, 1.3—3.0 mm thick, smooth, flaccid, and triangu-
lar in cross-section; 2—4 basal leaves per culm, 1—25 cm long, 1-4 mm wide,
flat, flaccid, and minutely scabrous toward the acute apex; 2 or 3 leaf-like
bracts subtend each inflorescence, varying in length, 2-25 cm long, 1—3(—
5) mm wide, spreading, flat, flaccid, and minutely scabrous toward the acute
apex; 6—24(—80) densely or loosely congested spikelets per spike; spikelets
8—18(—40) flowered, 3-8(-12) mm long, 1—1.5 mm wide, linear, and com-
pressed; rachillas 0.3 mm wide, 0.2 mm thick, dark brown, straight or slightly
curved, and wingless; scales (sometimes called “glumes”) subtending the
flowers 0.8—1.2 mm long, 0.8—1.5 mm wide, broader than long, 3-nerved,
tan or light brown at the center with narrowly hyaline margins, and the

Ww
bho
~

McKENZIE, ET AL., Cyperus fuscus in North America

Fic. 1. Cyperus fuscus. A. Habit (top and side views of clump and side view of two erect
plants). B. Inflorescence. C. Spikelet. D. Scale. E. Achene (including cross-sectional view
and side view of 3-cleft style). (Based on McKenzie 1802 with Jacobs, McKenzie 1804; McKenzie
1807 with Jacobs and Johnson, illustrated by Charles T. Bryson).

328 Sipa 18(1)

surfaces primarily dark reddish-brown to purple; the tips minutely apicu-
ate; achenes 0.9—1.1 mm long, 0.4—0.5 mm wide, gray to almost white,
trigonous, and saan ovoid; styles 0.4-0.6 mm long, 3-cleft, and not
persistent; stamens 2 per flower, 0.7—0.8 mm long; anthers 042055 aim

ong, tan or yellowish, and linear oblong.

—

_

BIOLOGY, ECOLOGY, AND DISTRIBUTION

On 9 September 1997, while conducting a search for state-listed species
of Cyperus, Schoenoplectus, and Lipocarpha along mudflats of the Missouri River
in Cooper County, McKenzie and Jacobs discovered a population of Cyperus
sp. unfamiliar to them. This Cyperus was semiprostrate with the culms ra-
diating like spokes in a wheel and leaning mostly horizontal to the ground
(Fig. 2). The most diagnostic features of the sedge were its dark purple to
reddish-brown scales, its strongly triangular stems, its bright rusty red roots,
its small spikelets, and its pale, trigonous achenes. Specimens were subse-
quently determined as Cyperms fuscus. McKenzie and Jacobs returned to the
site on 28 September 1997 and counted 110 plants scattered along the silty
mudflats and shoreline of the Missouri River, associated with the following
sens Ammania coccinea Rottb., Cyperus diandrus oa C. erythrorhizos Muhl.,
), odoratus L., C. squarrosus L. (C. aristatus Rottb.; C. inflexus Muhlenb.),
ae bioude (L.) L., Eragrostis hypnoides (Lam.) peor Sterns, & Pogg.,
Fimbristylis vablii Lam.) Link, Lewcospora multifida (Michx.) Nutt., Lipocarpha
micrantha (Vahl) G. Tucker, Polygonum spp. and Ranunculus scleratus L. Al-
though the majority of plants were in full sunlight, some extended into the
parcial shade of the developing seedlings of Sa/ix spp. and Populus deltoides
Bartram ex Marsha

Subsequent to this discovery, McKenzie and Jacobs were asked by Rhett
Johnson of the Missouri Department of Conservation to identify an unknown
specimen of Cyperus that had been collected from another site along the Missouri
River in adjacent Howard County, approximately 32 km WNW of the original
discovery site. McKenzie and Jacobs identified the specimen as C. fuscus
and visited the site with Johnson on 11 October 1997, where they counted
approximately 70 plants scattered along moist, sandy and silty flats adja-
cent toa chute that been formed during the 1995 flood of the Missouri River.

Because the discovery of C. fuscws in two adjacent counties along the Missouri
River suggested that the species could be wide

—

y distributed along the river,

Jacobs searched locations downstream of the original discovery site and discovered

it in Boone, Callaway, and Cole counties. A lack of time due to the season’s
first killing frosts prevented additional searches in other counties border-
ing the Missouri River. With the exception of Fimbristylis vahlii, associated
plants at the second, third, fourth, and fifth sites were mostly identical to
those previously mentioned for the original discovery site.

McKENZIE, ET AL., Cyperus fuscus in North America 329

G 2. Photograph of Cyperus fuscus taken from Lisbon Bottoms, Big Muddy National Fish
i. Wildlife Refuge, Howard Co., Missouri, 11 Oct 1997 @homeane by Paul M. McKenzie).

Although the initial North American discovery of C. fuscws in Massa-
chusetts was apparently along a sandy beach, many subsequent records along
the Atlantic seaboard were associated with “ballast sand” (Britton 1886),
or “waste ground, ballast, and wharf areas” (Rhoads & Klein 1993). The
Connecticut record comes from a “sandy river shore” (Tucker 1995) and habitat
for the species in California has been reported as “damp, disturbed soil, receding
shorelines, and puddles” (Tucker 1993). In the Midwest, the initial discov-
eries of this sedge were made along “wet open sandy flats” of the South Platte
River in Lincoln County, Nebraska and “wet sandy clay soil” along the banks
of the Keya Paha River in Tripp County, South Dakota (Weedon & Stephens
1969). The accounts by Britton (1886) and Rhoads and Klein (1993) sug-
gest that C. fuscus was initially introduced to the New England coast acci-
dentally via achenes that were in ship ballast water that originated in Eu-
rope. Introductions into other areas of the country, however, are more difficult
to assess. It is possible that waterfowl were responsible for the species’ in-
troduction into Nebraska, South Dakota, and elsewhere. Dunn and Knauer
(1975) postulated that waterfowl were responsible for the introductions of
Cyperus flavicomus Mich. {C. albomarginatus (C. Martius & Schrader ex Nees)
Steudel], Frmbristylis miliacea (L.) Vahl, and Aeschynomone indica L. into the
Mingo National Wildlife Refuge in southeastern Missouri.

330 Sipa 18(1)

While the source of introduction of C. fuscus into Missouri is unknown,
the recent and apparent widespread distribution of this sedge along the Missouri
River suggests that the species may have become established soon after the
1993 or 1995 floods. At the original discovery site in Cooper County, plants
were located at the spot where a large levee break had occurred in 1995.
Rolfsmeier (1995) reported that the two Nebraska records of C. fuscus were
from separate locations along the Platte River. Because the Platte River empties
into the Missouri River, and the Keya Paha eventually does the same, it 1s
feasible that Nebraska or South Dakota may have been the seed source for
the Missouri populations that became established along the Missouri River
following the 1993 and/or 1995 floods. This introduction may be due to
soil movement and sedimentation within the Missouri River floodplain.

Rice growers in the United States should be alerted to the potential of
this sedge becoming a troublesome weed. Holm et al. (1979) listed C. fuscus
as a principal weed in Portugal and as occurring in Afghanistan and Israel.
Based on the rapid and aggressive spread of its relative Cyperus difformis L.
in North American rice fields (Lipscomb 1980; Tyndall 1983; Bryson et al.
1996), and because C. fuscus is a rice weed in the Old World (Weedon &
Stephens 1969; Mingyuan & Dehu 1970; Holm et al. 1979), this species
should be looked for in rice producing areas of Arkansas, California, Loui-
siana, Mississippi, Missouri, Tennessee, and Texas.

Due to the combination of its semiprostrate habit with the culms radi-
ating like spokes in a wheel (Fig. 2), its dark purple to reddish-brown scales,
its strongly triangular stems, its bright rusty red roots, its small spikelets,
and its pale (almost white), trigonous achenes, this Cyperws is unlikely to be
confused with any other North American member of the genus. The con-
spicuous rusty red roots and small trigonous achenes are similar to C. erythrorbizos,
but the dark purplish scales, strongly triangular stems, apiculate achenes,
and stamen number (2 vs. 3 in C. erythrorhizos) easily distinguish it from
that species. The purplish-tinged scales of C. fuscus are somewhat reminis-
cent of the scales of C. diandrus, C. bipartitus Torrey, or C. flavicomus. It can
easily be separated from C. diandrus and C. bipartitus by its trigonous vs.
lenticular achenes, and by its smaller (ca. 1.0 mm vs. 1.8 mm) scales. It
differs from C. flavicomus by its flattened vs. erect habit, by the lack of white
margins on the scales, and by its trigonous vs. lenticular achenes.

he authors noted that bruised fresh and herbarium material of C. fuscus
possesses a fragrance similar to, but not as strong as, C. sguarrosus or C. setigerus
Torr. & Hook. Steyermark (1963) described ne odor as that of slippery
elm (U/mus rubra Muhl.). Bruhl (1995) stated that “(i)n a few genera of Cyperaceae,
a fenugreek (Trigonella foenum- graccum) odour is readily detectable in fresh
and (especially) herbarium material.”

There is apparently some disagreement in the literature regarding the

MCKENZIE, ET AL., Cyperus fuscus in North America a

number of stamens of each floret of C. fuscus. In the Flora of the Great Plains
(Kolstad 1986), two stamens are given in the key but three in the species
account. One (McGivney 1938), rarely two (Kiikenthal 1936), or two (Tutin
et al. 1980) stamens are also cited elsewhere in the literature. George Yatskievych
(Missouri Botanical Garden, pers. comm.) examined all voucher specimens
from Missouri and noted that florets had only two stamens.

Given the apparent rapid spread of this species along the Missouri River in
Missouri and the weedy nature of many Cyperus in the Old World, it is likely
that this species will be discovered with sufficient effort at other Midwest loca-
tions and possibly elsewhere in North America. While studying Cyperus speci-
mens at the Missouri Botanical Garden in 1988, Carter discovered a misidentifie
specimen of C. fwscus from western Nevada that originally was determined
as C. acuminatus Torr. & Hook. The “exposed mudflat” habitat at the Ne-
vada site is apparently similar to that in the San Joaquin Valley of adjacent
California described by Tucker (1993). Cyperus fuscus was not listed by Cronquist
et al. (1977) in the Intermountain Flora, nor in Kartesz’s (1987) dissertation
on the Flora of Nevada. As with California, Missouri, Nebraska, and South
Dakota, waterfowl may have been responsible for the introduction of achenes
of C. fuscus into Nevada. The oowine a puovide documentation for C,
fuscus in Missouri and Nevada with h ollowing Holmgren
et al. (1990), except ctb (pers. herb. of Charles T. Bae

Voucher specimens: U.S.A. Missouri. Cooper Co.: Big Muddy National Fish and Wildlife
Refuge-Overton Bottoms, ca. 1.9 mi NW of Overton, T49NR15 WSect.3 ISESENW 1/4,
Rocheport 7.5' Quad., 9 on 1997, McKenzie 1802 with Brad Jacobs (ctb, EIU, MO, SWSL,VSC);
ae 97-67 with Paul aa (MO); 28 Sep 1997, McKenzie 1804 with Brad Jacobs (ctb,

ard Co.: Big Muddy National Fish and Wildlife Refuge-Lisbon
see oms, ca. 1.5 mi emote tones 18, Arrow Rock 7.5' Quad., 11 Oct
1997, McKenzie oe with Brad Jacobs and Rhett Johnson (ctb, MICH, MO, SWSL, UMN,
UMO, VDB, VSC); Jacobs 97-40 with Paul McKenzie and Rhett Johnson (ctb, MO, MICH,
SWSL, UMN, UMO). Boone Co.: McBaine, W of Columbia waste water treatment la-
goon # 3, TA7NR13WSect. 7SWSESW 1/4, Hunstdale 7.5' Quad., 13 Oct 1997, Jacobs
97-37 with Tim Smith (MO, UMN, UMO). Callaway Co.: North Jefferson City, along N
side of the Missouri River, ee 400-500 m downstream from the Jefferson City
bridge over the Missouri River, T44N R11 WSect. 1SSWSW 1/4, Jefferson City 7.5' Quad.,
a Oct 1997, Jacobs 97-38 (ctb, MO, cy Jacobs 97-44 with Chris Dietrich (MO). Cole
o.: Jefferson = along the S edge of the Mi i River. imately 400-500 m downstream
ee the Jefferson City Bridge over the Missouri River, just W of Wears Creek, T44NR11WSect.
7NENE1/4, eae City 7.5' Quad., 15 Oct 1997, Jacobs 97-39 (MO, NEMO, UMO);
Jacobs 97-46 (MO). Nevada. Lyon Co.: Lahontan Reservoir, mudflats at the SE end of the
reservoir, T17NR26ESect.8, 4150 ft, plants growing on exposed mudflats, 1 Sep 1981,
Tiehm 6769 (MO).

ACKNOWLEDGMENTS

We greatly appreciate the assistance of George Yatskievych, Missouri
Department of Conservation, Flora of Missouri Project, Missouri Botanical

wa

325) Sipa 18(1

Garden, for initially identifying the Missouri specimens and providing additional
information on C.fuscus. We thank Connie Rutherford, U.S. Fish and Wildlife
Service, Ventura, California, for providing information on the species’ oc-
currence in California. Steve Rolfsmeier of Milford, Nebraska, graciously
supplied data on the Nebraska records. The helpful suggestions of George
Yatskievych and Robert Kral improved the quality of the manuscript.

REFERENCES

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new forms. Bull. Torrey Bot. Club 13:205—

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Bryson, C.T. and R. Carrer. 1992. Notes on Cine us and Kyllinga (Cyperaceae) in Missis-
sippi with on of six species new to the state. Sida 15:119—124,

Bryson, C.T. and R. Carrer. 1994. Notes on Carex, Cyperus, and Kyllinga ees in
Mississippi with records of eight species previously unreported to the state. Sida 16:17
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Bryson, C.T., J.R. MacDonatb, R. Carter, and $.D. Jones. 1996. Noteworthy Carex, Cyperus,
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Carr, B. 1988. Cyperus difformis L. (Cyperaceae) new to Texas. Sida 13 DIS 256.

CARTER, - L988. Cyperus hystricinus (Cyperaceae) new to Florida. Sida 13:118-119.

Carrer, R. and C.T. Bryson. 1991. A report of Cyperus grayioides and Cyperus ee
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SL:

Carter, R., C.T. Bryson, and = L. Lipscoms. 1987. Cyperus uniflorus (Cyperaceae) east of
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Carrer, R., R.L. Mears, K.C. a RKS, ond - T. Bryson. 1996. A report of four exotic Cyperus
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Jones, 8.D., A.A. Reznicek, and G.D. Jones. 1993. Cyperus setigerus (Cyperaceae), new for
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LipscoMB, B. L. 1978. Cyperus surinamensis (Cyperaceae): new to Arkansas, Kansas, and Oklahoma.
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Tucker, G.C. 1995. The vascular flora of southeastern Connecticut. Mem. Connecticut
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Rhodora 71:433.

THE USE OF ANIMAL-DISPERSED SEEDS AND
FRUITS IN FORENSIC BOTANY

BARNEY L. LIPSCOMB

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102, U.S.A. barney @ brit.org

GEORGE M. DIGGS, JR.
Department of Biology
Austin College, Sherman, TX 75090, U.S.A.
& Botanical Research Institute of Texas, ediggs@austinc.edu
ABSTRACT

A Specie case e of the peers use of animal- Sie aes propeguiess is s presented, and it is
1 h |

Cr utili ization DYT

id
Apna oe deeds and fruits are ene often cling tenaciously to clothes or other
materials worn or used by suspects, and are small and frequently go unnoticed. Further-
more, their identification is relatively inexpensive and tec niede Sean rd, and their
presentation as evidence is visually and intuitively obvious, making it ideal for the court-
room. It is also oe that forensic botany is an excellent topic to use as a case study in
pers botany or biology classes because of its inherent interest and integrative nature. In
order to facilitate such usage, a brief review of some aspects of forensic botany is presented
including references to pertinent literature.

RESUMEN

~ff5

Se presenta un ca p de propdgulos diseminados por animales en uso forense, y
se sugiere que este tipo ef evidencia puede tener mayor utilizaci6n en varios aspectos le-
ules. Las semillas y frutos dispersados por animales estén por todas partes, a menudo se

05

ae tenazmente a las ropas u otros materiales llevados o eee: por BOSD ese y
advertidos. ademas, su id

por ser p

barata y técnicamente sencilla, y I ia visual e intuitivamente,

convirtiéndose en ideal para los | ies Se sugiere cabies que la botdnica forense es un

tema excelente para ser usado como caso practico en las clases de sande por su interés
a bre de

algunos

inherente y naturaleza integrativa. Para facilitar ese uso se hace un
aspectos de la botdnica forense incluyendo las referencias bibliogréficas pertinentes.

The use of plants in justice and legal systems is thousands of years old,
probably beginning in such ways as trials by ordeal (Talalay et al. 1991;
Mabberley 1997). In these cases, suspects were forced to eat poisonous plants
and guilt or innocence was determined by survival. Presumably, this was
based on a psychological effect of guilt on the vomiting reflex—suppos-
edly, innocent individuals would expel the poisonous material, while the

Stipa 18(1): 335-346. 1998

336 Stipa 18(1)

guilty would retain the poison and thus die; the efficacy of such a tech-
nique is obviously questionable. The use of plant material as evidence has
also appeared in fictional works, such as the series of books by Ellis Peters
about the medieval Welsh monk/herbalist/sleuth, Brother Cadfael (W/hiteman
1995) and the works by H.C. Bailey about the detective, Reginald Fortune
(Bailey 1936, 1943).

The modern use of plants as evidence in a court case dates to the famous
1930s Lindbergh kidnapping case in which the son of Charles and Anne
Morrow Lindbergh was kidnapped and murdered. Largely through evidence
provided by botanist Arthur Koehler, Bruno Hauptmann was convicted of
the crime in 1935. Koehler demonstrated that the ladder used in the kid-
napping was built in part from wood taken from the attic of Hauptmann’s
residence (Tippo & Stern 1977; Baden 1983; Haag 1983; Lane et al. 1990;
Graham 1997). Koehler’s evidence included identification of the wood based
on anatomical characteristics, matching of annual growth rings, and unique
markings made on the wood by tools including a lumber yard planer and a
hand plane. Detailed accounts of the botanical evidence including photo-
graphs and graphics can be found in Tippo and Stern (1977), Haag (1983),
and Graham (1997). Graham (1997) gave an extensive list of references.
This was a landmark case, not only because it lead to the formation of fed-
eral kidnapping laws (Bock & Norris 1997), but also because the obvious
value of the evidence provided by Koehler set the stage for future forensic
uses of botanical information.

Since that time, there have been numerous other examples of forensic
botany (and other biological disciplines such as entomology—e.g., Rozen
and Eickwort 1997). The use of plant fragments, pollen grains or fungal
spores, plant trichomes (hairs), anatomical evidence from indigestible cell
wall material from the stomach contents of crime victims, molecular evi-
dence utilizing DNA, and ecological evidence useful in locating hidden graves,
crime sites, or dating when a crime occurred are just a few examples (Bock
et al. 1988; Lane et al. 1990; Mestel 1993; Yoon 1993; Blaney 1995; Bates
etal. 1997; Bock & Norris 1997; Graham 1997; Lewis 1997; Lindell 1997).
The following specific cases show some of the diversity in the field of bo-
tanical forensics. Lane et al. (1990) discussed a rape case in which leaves
and bark fragments were found in the pants cuffs of a suspect. The material
had gotten into his cuffs while he was climbing a tree to gain access to a
window of the victim’s house. His claim, that the victim had let him in
through a door, did not match the botanical evidence. Another example
used by Lane et al. (1990) involved a child abuse case. The parents claimed
that the child had been fed fruit cocktail just prior to dying. However, their
story was contradicted when his stomach contents showed no evidence of
the anatomically characteristic materials expected from such a meal (e.g.,

4

LipscomB AND Diccs, Forensic botany 37

stone cells from pears or needle-shaped crystals from pineapples). In the
case of a 1989 plane crash near Ruidoso, New Mexico, it was alleged that
an engine design flaw had allowed particulate matter (pollen) to build up
in the engine and cause the crash. However, it was shown that since the
pollen was in fresh condition and had normal cytoplasm and cell walls as
seen by electron microscopy, it could not have been exposed to the high
temperatures present during engine operation or in the post-crash fire that
distorted even aluminum. Further, the pollen was that of insect-pollinated
plants found near the storage site of the plane wreckage; such pollen grains
are unlikely to be found in any quantity in the atmosphere. Based on the
evidence from forensic palynology, it was concluded that the pollen had gotten
into the wreckage post-crash during several months of storage and there-
fore had nothing to do with the accident (Blaney 1995; Brunk 1997; Gra-
ham 1997; Lewis 1997). A final example is the use of molecular evidence
linking a murder suspect toa palo verde tree (Parkinsonia aculeata L., Fabaceae)
at an Arizona crime site where he allegedly Auimaped the body ofa victim.
Plant geneticist Tim Helentjaris of the University of Arizona demonstrated
that two seed pods found in the suspect’s truck came from a specific palo
verde tree scraped by the suspect’s truck at the crime scene. This example is
important because it was the first in which plant DNA was used in a crimi-
nal case (Mestel 1993; Yoon 1993). Overviews of forensic botany were pro-
vided by Lane et al. (1990) in the general science literature and by Bock
and Norris (1997) in the forensic science literature.

Our interest in this topic has developed over the course of a number of
years during which as plant taxonomists we have been called upon numer-
ous times by poison centers, hospitals, and law enforcement agencies to identify
plants or their fragments. We agree with Bock and Norris (1997) that fo-
rensic botany is a resource underutilized by the law enforcement commu-
nity. Further, we believe that forensic botany can be very effectively used in
botany or biology courses to show the importance, applicability, and inte-
grative nature of botany. Because forensic botany cuts across all botanical
disciplines and because a given case may require many research approaches
and techniques, it is a discipline that can stress the integrative nature of
botany and science as a whole. Additionally, it is an excellent topic to use in
emphasizing problem solving and critical thinking skills. The purpose of
the present article is thus two-fold. First, based on our successful use of such

information in college teaching, we want to provide in an easily accessible
botanical journal a brief review of forensic botany and appropriate refer-
ences in order to encourage further such usage. Second, we present a spe-
cific case of the forensic use of animal-dispersed propagules and suggest that
this type of evidence has the potential to be more widely used in criminal
investigations.

338 Sipa 18(1)

On July 12, 1995, a sleeping two year old girl was pulled from the first
floor window of an apartment in Fort Worth, Texas. The child was sexually
molested, but fortunately was found alive about three hours later in a weedy
area several hundred meters from where she was abducted. Assorted evi-
dence was used in the case including fingerprints and DNA from semen.
However, the easily understandable botanical evidence was an important
factor in convincing the jury of the suspect’s guilt. Because the Botanical
Research Institute of Texas (BRIT) is a well known source of botanical in-
formation in the local community, we were contacted by the district attorney's
office to identify tiny plant fragments taken from the shoelaces of the sus-
pect. In addition, we were provided with a bag of assorted plant material
that had been collected from the crime scene where the child was left (Fig.
1). Upon opening the evidence envelope, we immediately recognized the
ca. 4 mm long plant fragments as single-seeded mericarps from a member
of the Apiaceae (carrot family). Using a dissecting scope and authenticated
specimens in the Botanical Research Institute of Texas herbarium, these were
identified as mericarps of Tori/is arvensis (Huds.) Link, an introduced spe-
cies commonly known as hedge parsley. The bag of material from the crime
scene was then examined and a mature, fruit-bearing plant of hedge pars-
ley was found. Under a dissecting scope, the small mericarps of this species
(Fig. 2) are very distinctive. They are densely covered with bristles tipped
with microscopic barbs that enhance their attachment to fur or clothing.
They also have several very characteristic lines of closely appressed hairs between
the bristles. Large photographs of mericarps from both the suspect's shoe-
laces and the crime scene (Fig. 3) were presented in court by one of us (BLL).
Like fingerprints, this was distinctive visual evidence, more easily under-
stood than the scientifically sound but conceptually complex evidence pro-
vided by molecular techniques such as DNA analysis. Because the suspect
could have possibly picked up the mericarps from some other location, the
botanical evidence alone would not have been sufficient for a conviction.
However, it firmly linked the suspect with the crime site and in combina-
tion with other evidence was successfully utilized by prosecutors Sharon
McLauchlin and Larry Thompson and criminal investigator Dennis Timmons.
The suspect, David Noel Saddlemire, was convicted of aggravated eae a
ping with the jury taking only 55 minutes to reach their verdict (Fig. 4).
He was subsequently sentenced to 99 years in the Texas state prison system.

Ectozoochory, the transport of a diaspore or propagule on the outside of
an animal, is a common mechanism of dispersal among flowering plants
(van der Pjil 1982). While there are various types of diaspores (e.g., vegeta-
tive bulbils, fragments of the parent plant), the most common types are
seeds, whole fruits, or fruit segments (e.g., mericarps as in the above ex-
ample). Various methods of attachment are known, ranging from viscous

LirscomB AND Diaccs, Forensic botany oie)

FORT WORTH POLICE DEPARTMENT
E LABORATORY

RECEIPT

CASE NO. 141868

TYPE CASE Kidnapping/Sexual Assault Child

OFFENSE NO. 95 364266

VICTIM A. S.

SUSPECT David Saddlemire

Raceived from “Hoary ey [ii pSCom la fen
iS)
Title and Address: Ash, Dice. Rat TOF See FA barth Foor

DATE T: $Ib TIME 1220 wom
w_ PK IT

DESCRIPTION

Received botanical evidence as listed below:

A tape sealed brown paper bag containing:

1. A tape sealed bag holding plant material
(ll - Invoice #95 See

material ee from right

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ao
©
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subsamp ant materia
(28 - pene #95C03471
3. <A tape sealed g holding plant aubeeiet
(28 - Invoice Ear
4. A Sra ood

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drrcdeye oars wan Vardy
aN Lonernn fpr SIGNED. TT
whognoplnsy pass CUD one eS
TELEPHONE NO. 1) BTV SOKY

001-35506-025
8-1-77

Fic. 1. Receipt of botanical evidence from the Fort Worth Police Department Crime
Laboratory.

340 Sipa 18(1)

Fic. 2. Torilis arvensis. A) habit; B) flower; C) fruit showing mericarps with uncinate bristles;
D) cross-section of fruit (drawn by Linny Heagy).

LirscoMB AND Diccs, Forensic botany 341

LeftShoe Right Shoe

Known

<---]| Cm--->

Fic. 3. Photographs of mericarps from both the suspect’s shoelaces and the crime scene
ii es by Larry A. Reynolds, courtesy of Tarrant County Medical Examiner).

or mucilaginous substances that cause the propagule to stick to the dis-
persal agent, to very sharp, barbed or recurved hooks, spines, or awns (van
der Pyjil 1982). We believe that many of these examples are potentially valuable
to law enforcement agencies. From many field trips both for research and
with students, it has been our experience that it is rare to return from the
field without attached plant material either on the clothes, in pant cuffs,
on socks, or embedded in shoelaces. Some of these are merely annoying,
while others (e.g., Arzstida species—threeawn grasses) are extremely irri-
tating when penetrating socks or pants. Many of these seeds or fruits fre-
quently get transferred to the interior of cars on carpeting or upholstery
One does not have to go far from the sidewalk to encounter such plant materials.
Because many abundant weedy species are animal-dispersed, yards, aban-
doned lots, and virtually any weedy or disturbed site will have some such
species. Because these plant materials are so frequently encountered and because
different plant species are found predominantly in specific habitats and during
particular seasons, they are potentially valuable sources of evidence that can
link suspects with crime scenes both spatially and temporally. Also, many
(e.g., tiny fruit segments of some Apiaceae or tiny fruits of some grasses)

342 Stpa 18(1)

TARRANT COUNTY
OFFICE OF THE
CRIMINAL DISTRICT ATTORNEY
TIM CURRY JUSTICE CENTER
CRIMINAL DISTRICT ATTORNEY 401 W. BELKNAP
817/884-1400 FORT WORTH, TX 76196-0201

April 21, 1997

Mr. Barney Lipscomb

Botanical Research Institute of Texas
509 Pecan Street

Fort Worth, Texas 76102

Dear Mr. Lipscomb:
ale su for your expertise and assistance in the David Saddlemire trial. The case was

our testimony was le helpful in explaining to the jury one of the
eirchmnsinnwes surrounding the offen

he defendant was found guilty and sent d to 99 he p Again,
thank you for your assistance. This case was an important to the State of Texas and this
community.
Sincerely,

if, ; iy
Peer J Me ut aN
Sharon McLauc ide

Assistant Criminal District Attorney

Fic. 4. Letter from Tarrant County Office of the Criminal District Attorney confirming
cone jury conviction of David Noel Saddlemire.

become deeply imbedded in cloth or carpeting, go virtually unnoticed, and
often remain attached even after repeated washings or other types of clean-
ing. Further, seeds and fruits are easily and inexpensively identified by trained
botanists using nothing more than a 10X hand lens or inexpensive dissect-
ing scope, taxonomic literature, appropriate illustrations, and herbarium
specimens. While molecular forensic techniques can potentially provide
very definitive information, they are much more expensive, require sophis-
ticated laboratory facilities, and are less intuitively obvious for courtroom
presentation.

>

LipscoMB AND Diccs, Forensic botany 343

Sometimes the attachment of ectozoochorous propagules can be quite
tenacious. This means that they could be attached to a perpetrator’s cloth-
ing for a very long time, or even enter the flesh. The following are specific
examples from mammals that illustrate the point. Sharp-pointed structures
such as fruits or awns can penetrate the mouth or other tissues of livestock
(and have to be extracted by pliers) or even become subcutaneous and re-
quire surgical removal. Veterinarians (John Brakebill, Larry Edwards, Ken
Lawrence, pers. comm.) indicate that it is not uncommon to find grassburs
(Cenchrus species), awns (e.g., from Hordeum species—foxtail) or the pointed
fruits of needle, spear, or threeawn grasses (Nasse//a or Aristida species) in
animals. For example, in North Central Texas, Cenchrus burs are often found
embedded between the toes of dogs and grass fruits are known to penetrate
buccal tissues including the tongue causing serious problems (Ken Lawrence,
pers. comm.). Perhaps more striking is the ability of the sharp fruits of needle
grass (apparently, Nassella leucotricha (Trin. & Rupr.) Barkworth [formerly
known as Stipa leucotricha Trin. & Rupr.}—commonly called winter grass,
Texas winter grass, spear grass, or Texas needle grass) to deeply penetrate
flesh. These can become subcutaneous and require surgical removal; for example,
they can enter between the toes of dogs and sometimes migrate long dis-
tances subcutaneously causing draining tracts that will not heal until the
fruit is removed (Larry Edwards, pers. comm.). The most extreme case we
are aware of involved a fruit that penetrated through the skin and then the
chest wall of a dog, eventually becoming embedded in a lung and causing
a fatal case of pneumonia (John Brakebill, pers. comm.). Also tenacious are
the spiny fruits of Trzbu/us terrestris L., puncture vine, of the Zygophyllaceae.
These are very painful to both animal and human feet, damage even tires,
and are occasionally fatal to livestock if eaten (Correll & Johnston 1970); it
would not be surprising to find them attached to various objects and pos-
sibly even the tires of a suspect’s vehicle.

Locally in North Central Texas, we believe So/iva pterosperma (Juss.) Less.,
lawn burweed, (Asteraceae) collected from a soccer field near Arlington, Tarrant
Co. (1995), was possibly spread from eastern Texas by athletic shoes; its
fruit is easily, and painfully, attached by its persistent, spine-like style (Diggs
et al., forthcoming). Such propagules could remain attached to a suspect's
clothing or shoes for long periods of time. Other Asteraceae are well-known
as being animal-dispersed with the pappus of many species being modified
into an attachment structure. The retrorsely barbed awns of Bidens species,
beggar’s ticks, are strikingly effective. In another composite genus, Xanuthium,
cocklebur, the surface of the bur is conspicuously covered with stiff, hooked
prickles ca. 5 mm long and the bur is also terminated by two prominent
spines. The attachment of the hooked prickles to clothing or shoelaces is

344 Stipa 18(1)

very effective and they can also easily penetrate human skin. The hooks are
strikingly reminiscent of velcro. According to the VELCRO® Industries
homepage (www.velcro.com), in the early 1940s, a Swiss inventor, George
de Mestral, after a walk noticed “cockleburrs” [presumably Xanthinm] on
his dog and his pants. He examined the hooked prickles under a micro-
scope and derived the idea for the well known two-sided fastener—one side
with stiff, cocklebur-like “hooks” and the other side with soft “loops” like
the cloth of his pants. The word velcro comes from the French words ve-
fours, velvet, and croché, hooked.

While some of the examples above were presented to show the tenacity
with which diaspores attach, commonly the seeds or fruits are small and
merely cling to the fur, feathers, feet, beak, etc. of the dispersal agent with
little or no adverse effect. Because they are often small and inconspicuous,
they may be particularly valuable from the forensic standpoint. A well known
such member of the Fabaceae (legume family) is the genus Desmodium, tickseeds.
The fruits or loments are jointed and break apart into 1-seeded flat seg-
ments that are the dispersal units; they easily attach to hair or clothing. In
North Central Texas for example, there are 12 members of this genus, a
number of them occupying rather specific habitats (Diggs et al., forthcom-
ing). Other well known examples are the numerous Apiaceae that have small
schizocarps (a fruit that splits between carpels into one-seeded portions called
mericarps) whose mericarps are covered with bristles or hairs and become
readily attached to many surfaces. These are particularly well known to owners
of long-haired dogs because large numbers of the mericarps become entangled
in the fur—sometimes the situation is so severe that the only recourse is to
shave the dog. The final example given here is the legume genus Medicago,
commonly known as bur-clovers. There are numerous introduced species
of this genus, many of which have prickly fruits. These fruits can be some-
what larger than those mentioned previously, but are still effective at at-
taching to dispersal agents. Numerous other examples could be given which
have potential use forensically. Fortunately, most are easily identified by
experienced field botanists because such researchers have encountered them
many times on their own clothing or equipment.

Other less obvious diaspores could also be potentially useful. For example,
at the present time, several invasive aquatic species (e.g., Hydrilla verticillata
(L.f.) Royle, in the Hydrocharitaceae) are spreading in North Central Texas,
apparently by power boats transporting vegetative propagules (plant frag-
ments). In areas of the country where there are numerous relatively new
reservoirs and where the distributions of many aquatics, especially intro-
duced species, are spotty, plant material of a given species could easily be
used in linking a suspect with a given body of water.

ws)
os
A

LipscoMB AND Diccs, Forensic botany
SUMMARY

Forensic botany is a developing discipline that potentially has broader
applicability than is seen at present. Technically simple, visually obvious,
easily understood, and inexpensive methods such as the use of animal-dis-
persed seeds and fruits are particularly worthy of further consideration. Because
of the inherent interest in criminal cases, the potential for emphasizing problem
solving and critical thinking skills, and the integrative nature of the sub-
ject, forensic botany is a field that can be useful in botany and biology edu-
cation.

ACKNOWLEDGMENTS

We would like to thank our colleagues at various law enforcement agen-
cies, poison centers, and hospitals for their cooperation and hard work. Special
thanks to Dennis Timmons, Criminal Investigator of the Tarrant County
Office of the Criminal District Attorney, and Larry A. Reynolds, Forensic
Photographer, for the photographs, which are used courtesy of the Tarrant
County Medical Examiner. Thanks also to John Brakebill, D.V.M., Larry
Edwards, D.V.M., and Ken Lawrence, D.V.M., veterinarians in Sherman,
Texas, John Lanzalotti, M.D. and Bill Vande Water, BSI, for information on
fictional uses of plants as evidence, and Kenna Pirkle, an Austin College stu-
dent, whose Plant Biology class paper on forensic botany prompted us to
write this manuscript. Linny Heagy provided the illustration of Tori/7s arvensis.

REFERENCES
BabeNn, M.M. 1983. Plenary session: The a kidnapping revisited: Forensic sci-
ences then and now. J. Forensic Sci. 8:1035—
Baitey, H.C. 1936. A clue for Mr. Fortune: ae at his best. The Sun Dial Press, Garc
City, NY.

jon

en

. 1943. The thistle down. Ellery Queen’s Mystery Magazine 4:5-15.
Bates, D.M., G.J. to and R.D. Lee. 1997. Forensic botany: Trichome evidence. J.
Forensic Sci. 42:380—
Bianey, C. 1995. Who y ya gonna call? Bioscience 45:746-747.
Bock, J.H., M. Lang, and D. Norris. 1988. Identifying plant food cells in gastric contents
for use in forensic investigations

a laboratory manual. U.S. Department of Justice, National
Institute of Justice, Washington,

Bock, J.H. and D.O. Norris. 1997. Forensic botany: An under-utilized resource. J. Foren-
sic Sci. 42:364-367

Brunk, 8.K. 1997. The Ruidoso plane crash—the background and the trial verdict. J. Forensic
Sci. 42:378-379

CorrELl, D.S. and M.C. JOHNsToNn. 1970. Manual of the vascular plants of Texas. Texas
Research Foundation, Renner.

Diccs, G.M. Jr., B.L. Lipscoms, and R.J. O'KENNON. Shinners & Mahler’s illustrated plants
of North Central Texas. To be published in late 1998 by the Botanical Research Insti-
tute of Texas, Fort Worth.

346 Stipa 18(1)

GraHaM, A. 1997. Forensic palynology and the Ruidoso, New Mexico plane crash—the
pollen evidence II. J. Forensic Science 42:391—

GranaM, S.A. 1997. Anatomy of the Lindbergh kidnapping. J. Forensic Sci. 42:368—377.

Haac, L.C. 1983. The investigation of the Lindbergh kidnapping case. J. Forensic Sci.
8: 1044 1048.

Hornan, J.J. 1983. The investigation of the Lindbergh kidnapping case. J. Forensic Sci.
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Lange, M.A., L.C. ANDERSON, T.M. Barkiry, J.H. Bock, E.M. Girrorp, D.W. Hatt, D.O.
Norris, T.L. Rost, and W.L. STERN. 1990. Forensic a Bioscience 40:34—39.

Lewis, W.H. 1997. Pollen composition ina crashed plane’s engine. J. Forensic Sci. 42:387—
200.

Qu.

LINDELL, C.M. 1997. Field sampling and chemical analysis. J. Forensic Sci. 42:398—400.
MasBer ey, D.J. 1987. The plant book, a portable dictionary of the higher plants. eee
with corrections in 1989. Cambridge Univ. Press, Cambridge, United Kingc

YS

1997. The plant book, a portable nena of the higher ci “Ond ed,
Giehiides Univ. Press, Cambridge, Culeed Kingdo
Mestet, R. 1993. Murder trial features tree’s genetic ake New Scientist 138:6.
ROZEN, J.G. and G.C. Eickwort. 1997. The entomological evidence. J. Forensic Sci.
42: ie fa
TALALAJ, Ks Ta LALAJ, and J. TALALA). 1991. The strangest plants in the world. Hill of
ee Publishiee Co., Melbourne, Aust ralia
Tippo, O. and W.L. STERN. 1977. Humanistic Bore: W.W. Norton & Co., New York.
VAN Der Py, L. 1982. Principles of dispersal in higher plants. Springer-Verlag, Berlin.
WHITEMAN, R. 1995. The Cadfael companion: The world of Brother Cadfael. Mysterious
Press Books, New Yor
Yoon, C.K. 1993. Daccouedl witness for the prosecution. Science 260:894—-895.

NOTES

GLAUCIUM CORNICULATUM (PAPAVERACEAE) IN TEXAS

In May 1993, the senior author was informed by her friends, Don and Jo
Robison, about an unusual population of wildflowers growing on their ranch.
The population has been growing in the same field since the early 1960s
when it first appeared with a cultivated batch of cotton seed (D. Robison,
pers. comm.). The cotton was eventually replaced with sorghum alum and
other grasses in order to graze cattle and the plant has continued to remain
despite grazing upon by cattle during periods of severe drought.

Thousands of these individuals were observed by the senior author growing
in a sandy field. The majority of the plants, however, were toward the end
of their blooming period and had already developed their fruits (long lin-
ear capsules full of tiny black seeds). The senior author took the plant to be
a poppy, however, one that she was unfamiliar with.

This plant was later identified as Glaucium corniculatum (L.) Rudolph by
the junior author by comparison with herbarium collections at TEX. For-
tunately, there was already a collection (O’Kennon & Cheatham s.n.) of Glauctum
from Texas. This specimen, however, was incorrectly identified (as G. flavum,
a perennial species with yellow petals and glabrous capsules).

In the years since 1993, Texas experienced a drought and few, if any, of
the Glaucium in the Garza population had bloomed, preventing further
observations by the senior author. In April 1997 sufficient rains brought
the Glaucium population to bloom allowing the senior author to both ob-
serve and photograph (Fig. 1) the population. Later, perhaps as a result of
the rains, a third population of G/awcium was brought to the attention of
the junior author (Bill Henderson, pers. comm.). This population was col-
lected at the Balcones Canyonlands National Refuge. An estimated 100 plants
were observed growing ina circular area of about 30 feet in diameter. Two
individuals were blooming and five were in fruit, but the majority of the
plants were in a juvenile phase too young to flower.

The genus Glaucium is native to Southern Europe, the Mediterranean,
Hungary and South Central Russia. It has become adventive in California,
Kansas, Montana, Nevada, New York, Oregon and Pennsylvania (Barkley
1986; Hickman 1993; Kiger 1997). It is immediately distinguished from
other Texas genera in the Papaveraceae by its fruits which are straight, unribbed,
linear, capsules up to 25 cm long. Glaucinm corniculatum is a wholly pubes-
cent annual, 30—40 cm tall, caulescent, with clear yellow sap, cauline deeply
pinnate unprickled leaves, inflorescence of a solitary flower that has two
free sepals and four petals. The petals are a vivid scarlet color, two of them

Sioa 18(1): 347. 1998

348 Sipa 18(1)

are wider than the other but all are of equal length (3—4 cm). At the base of
each petal is a purple pattern that extends toward the tip about 1/3 the
length. The pattern is oval, 1O mm wide and lined with a creamy feather-
like design (Fig. 1).

Although collected in Texas as early as 1986 (Keeney 5906) and recently
reported in Texas (Jones et al. 1997), and North America (Kiger 1997),
Glaucinm corniculatum has 4 to be officially documented in the state. Pre-
sented here is an updated key to the genera of Papaveraceae in Texas, a list-
ing of known voucher specimens, and a photo of the flower (Fig. 1).

Below is a revised key to the genera of Papaveraceae in the Manual of the
vascular plants of Texas (Correll & Johnston 1970).

1. Acaulescent herbs; petals 8 or more, not crumpled in the bud; stigmas over

the valves of the capsule. Sanguinaria

. Caulescent herbs or (of subacaulescent) confined to the Trans-Pecos; petals

4 to 6, mostly crumpled in the bud; stigmas over the placentae 2
2. Perianth and stamens borne on the rim of the hypanthiumlike expansion
of the receptacle; sepals united into a calyptra; fruit conspicuously ribbed.
Eschscholzia
2. Perianth and stamens strictly hypogynous; sepals not united into a calyp-
tra; fruit not ribbed. 3
3. Herbage prickly; flowers on short pedicels; capsule dehiscing from the
apex by 4 to 6 valves. Argemone
3. Herbage not prickly; flowers on long pedicels 4
4. Stigmatic disk present; capsules short and turgid, to 5 cm long,
dehiscing by means of small openings just beneath the truncate
summit. Papaver
4. Stigmatic disk absent; capsules linear up to 25 cm long, dehisc-
ing longitudinally from the apex. Glaucium

Glaucium corniculatum (L.) Rudolph, Fl. Jen., 13.1781. Chelidoninm corniculatum
L., Sp. Pl. 1:56. 1753.

Voucher specimens. TEX AS. Garza Co.: community of Justiceburg, on Farm Rd. 3519
on the way to Lake Alan Henry, Don and Jo Robison Ranch, SE corner of Garza Co., ca. 3
mi from Kent Co. line, in field on N side of dirt road, 10 May 1993, Kirkpatrick s.n. Gao.
Kerr Co.: W of Hunt on TX 39, dry creek bed on N side of road, 2 May 1993, O’Kennon &
Cheatham s.n. (VEX). Travis Co.: Balcones C eye National eae 0.2 mi c.

dirt road NE ate intersection with Cow Creek road, dirt road is 3.4 N along Cov
Creek Road fr¢ 31, just after large yellow house (30° 33' 50" N, an O7' 70" a
May 1997, /. K. ee as 150 (TEX). San Saba Co.: along the aon Cre

William Clark farm at Bend, on the edge of an over grazed field, May 1986, Keeney
5906 eT); SE of the William Clark house on the east side of je Cherokee Creek at
Bend (1654 plants present), 9 May 1987, Keeney 6642 (BRIT).

The following partial list of species from the Travis county population

Sipa 18(1): 348. 1998

Notes 349

Fic. 1. Photo of Glaucium corniculatum (L.) J.H. Rudolph. Photo by Z. Kirkpatrick.

emphasizes the weedy nature of the vegetation at this site: Argemone albiflora
Hornem., Buglossoides arvense (L.) I.M. Johnston*, Callirhoe involucrata (Tort.)
A. Gray, Centaurea melitensis L.*, Cirsium texanum Buckl., Convolvulus equitans
Benth., Cucurbita foetidissima Kunth, Evax sp., Gaura calcicola Raven & Gregory,
Glandularia bipinnatifida (Nutt.) Umber, Laminm amplexicaule L.*, Lepidium
virginicum L., Marrubinm vulgare L.*, Medicago lupulina L.*, Hill., Oenothera
ead Nure: nis salis cinavescens (Doe) A. Hitch. Plane ph acanntie
Dene., Rapistrum rugosum (L.) ee (Sims.) D. Don., Sa/via
seh aiken Benth. See haler Small, = naturalized species.

—Zoe Raboahie Box 696, Post, TX 79356, U.S.A. and J.K. Williams, De-
partment of Botany, University of Texas, Austin, TX 78713, U.S.A

REFERENCES
BarkLey, T.M., ed. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence.
Corre.t, D.S. and M.C. JoHNsron. 1970. Manual of the vascular plants of Texas. Texas
Research ee Renner, Texas.
Hickman, J.C., 1993. The Jepson manual: Higher plants of California. University of
California = Berkeley.
Jonss, S., J. Wie aia PM. Coe ry, 1997. Vascular plants of Texas: A comprehensive

—

checklist including sy y, bibliography, and index. University of Texas Press, Austin.
Kicer, R.W. 1997. 7 Glan tuCLUM?. o oe Nc ig penesica Editorial Committee, eds. Flora of
North America: Volume 3. M j ‘ 1 Hamamelidae. Oxford University

Press, Oxford. Pp. 302— 304.

Stipa 18(1): 349. 1998

350 Sipa 18(1)

HOVENIA DULCIS (RHAMNACEAE) NATURALIZED
IN CENTRAL TEXAS

In the late summer of 1997 an unusual tree about 12 m tall was observed at
the base of a high limestone cliff above Town Lake on the Colorado River in
Austin. It had alternate leaves with prominent veins, and appeared to have
strange fruiting structures high on the branches, these structures apparently
deformed with galls. The tree was identified as a member of the Rhamnaceae
based on the characteristic three-carpellate fruit with a remnant disk at the
base, very similar to Co/ubrina. Comparison with the Rhamnaceae collec-
tion at TEX revealed the tree to be Hovenia dulcis Thunberg (Fig. 1), the
Chinese raisin tree, which is known to be cultivated in Texas (Jones et al.
1997). Mabberley (1997) suggested Hovenia to be of close affinity to Colubrina,
while recent work by Richardson (unpublished data) suggest that it 1s more
closely-allied with Zzzzphus. What appeared to be galls are actually peduncles
that swell naturally as the fruit develops.

This species is native to mesic forests in China and was introduced into
India and Japan for its edible peduncles (Hooker 1872; Rehder 1940; Roxburgh
1975; Sargent 1916). In China the swollen peduncles of the cymose inflo-
rescences have long been used medicinally “to offset the effects of over-in-

dulgence in wine” (Sargent 1916). The peduncles are especially palatable
after frost when they redden and the juice sweetens with pear-like flavor
(Mabberly 1990; Reich 1991). The fruit itself is not edible.

This tree is known for its cold-hardiness and is cultivated in USDA cli-
mate zone 5, with minimum temperatures of -20°F (Dirr 1990; Staff of the
L.H. Bailey Hortorium 1976), the equivalent of Milwaukee, WT or Binghamton,
NY. It is therefore surprising to find it naturalized in Austin, TX, an area
that is much warmer, and more importantly, significantly drier than much
of zone 5. The locality where it grows in Austin is a cool, moist microcli-
mate, at the base of a cliff of Edwards limestone and dolomite perhaps 40
m high, facing directly to the northeast. This cliff reaches almost to lake
level, except for a very short but steep talus slope of boulders and loamy
soil. Vegetation growing at the base of this northeast-facing cliff is almost
completely protected from mid-day and afternoon sun, and the presence of
a body of water surely modifies the microclimate further. Further explora-
tion via boat several weeks later resulted in the discovery of a larger tree
perhaps 100 m upstream, and a smaller tree perhaps 100 m downstream,
both at the base of the same cliff. Flowering specimens were later collected
in May 1998 from the first tree discovered.

It is possible that this small population of H. dulcis is descended from
cultivated specimens that existed at the University of Texas College of Pharmacy

Sipa 18(1): 350. 1998

Notes 451

¥

erent

Fic. 1. Fruiting branch of Hovenia dulcis Thunb., showing capsules and swollen peduncles.
Bar = 1 cm

Drug Garden, the only other specimens known from the Austin area. Prior
to the mid-1940s this garden existed about four miles to the north-east of
the collection site, and was subsequently abandoned and never reconstructed
(Beryl Simpson, pers. comm.). Two specimens exist at TEX from this gar-
den, dated 1940 and 1946. It is possible that while H. du/cis was cultivated
in this garden, birds ate the fruit, roosted on the cliffs and trees by the river,
and defecated the seed to the forest soil far below. Until the early 1960's
this portion of the Colorado river was still free-flowing, and large stretches
of loamier soil, which is preferred by H. duw/cis (Dirr 1990; Staff of the L.H.
Bailey Hortorium 1976), likely existed. When Town Lake was created in
the early 1960’s, perhaps a much larger population of this species was sub-
merged and destroyed, leaving only a small number of individuals grow-
ing near the top of the appropriate habitat. During further explorations, no
seedlings were observed, and seedling success is probably quite low because
numerous other species compete for space on this very thin ribbon of land.
Furthermore, the majority of fruiting branches overhang the water, so most

SIDA 18(1): 351. 1998

352 Sipa 18(1)

fruit falls directly to the bottom of the lake. Similar habitats exist within

the region, so it is possible that H. di/cis occurs unobserved elsewhere.

Voucher specimens: TEXAS. Travis Co.: tree 40 ft tall, growing at the base of a high,

NE-facing limestone cliff along the edge of Town Lake, with Cornus drummondii, Platanus

occidentalis, and Smilax bona-nox, across the lake from the boat ramp at the University of
nridge Field Lab, Austin, 30 Sep 1997, D. Goldman with J. Crutchfield 1105

(BH, BRIT TEX): same locality but with flowers, 29 May 1998, D. Goldman with P. Griffith

ae

— eee BRIT, TE
. Travis Co.: cultivated, Austin, 16 Sep 1940, C.C.

her ined: TEXAS
a s, (TEX); ex cultivation; Drug Garden, University of Texas, 24 May 1946, C.C.
Albers 46pb060 (TEX).

Thanks is given to John Crutchfield and Patrick Griffith for their assis-
tance in obtaining specimens of this plant, and Hobbes Goldman, Beryl
Simpson, Billie Turner, Justin Williams, and Tom Wendc for their-assis-
tance with the manuscript.—Donglas . Goldman, Department of Botany, University
of Texas, Austin, TX 78713-7640, U.S.A.

REFERENCES

— M.A. 1990. Manual of woody ce peal Their identification, ornamental
ropagation and use Stipes Publishing, Champaign.

ACTELISTICS cultur re p
Hooker, J.D. 1872. The flora of British india, part 1. L. Reeve & CO., London.
Jones, $.D., J.K. Wiper, and P.M. Montcomery. 1997. Vascular plants a Texas. A com-

ptehessive checklist. U niversity of Texas Press, Austin.
Maspertey, D.J. 1997. The plant book. Second edition. Cambridge University Press, Cambridge.
Reuper, A. 1940. Manual of cultivated trees and shrubs. Macmillan Publishing Co., New

York.
Reicu, L. 1991. Uncommon fruits worthy of attention. Addison-Wesley, Reading.
RoxspurGH, W. 1824. Flora Indica. Reprinted by Oriole Editions, 1975, New York.
SARGENT, C. 8. 1916. Plantae Wilsonianae, vol 2. Cambridge University Press, Cambridge.
Stare OF THE L.H. Batley Horrorium. 1976. Hortus third. Macmillan Publishing Co., New
York.

Sipa 18(1): - 1998

Notes 353

HABRANTHUS TUBISPATHUS (LILIACEAE) NEW
TO THE FLORA OF ALABAMA

The Copper Lily Habranthus tubispathus (LU Her.) Traub was first collected in
Alabama in che northeast part of the city of Troy in Pike County on 15 July
1988 during field work for a Flora of Pike County. This species was then
collected in the community of China Grove in the northern part of Pike
County on 17 July 1995 and in rural western Bullock County near the area
referred to as High Ridge on 12 July 1996. A fourth site southwest of the
city of Troy in Pike County was discovered in August of 1997 (Fig. 1). These
are the first records of this species from Alabama, and the second record of
this species in North America outside of Texas and Louisiana according to
Holmes and Wells (1980) and Burkhalter (1984).

Habranthus tubispathus (L’Her.) Traub (Liliaceae), also known as Zephyranthes
texana Herbert or Habranthus texanus (Herbert) Herbert ex Steud., is a sca-
pose perennial from a small bulb. Each 10-20 cm scape bears a single yel-
lowish-orange flower produced during the summer months, usually following
a rain. The leaves are linear, 3-5 mm wide and up to 25 cm in length, and
appear in aucumn and wither by early spring. The fruit is ovoid and three
lobed containing numerous flat, black papery seed at maturity. The genus
Habranthus is separated from the genus Zephyranthes by Sealy (1937) based
upon the filaments of Habranthus being of four lengths and the anthers af-
fixed at the middle, while in Zephyranthes the filaments are of alternate lengths
and the anthers affixed below the middle.

In the city of Troy, Copper Lily is found growing in lawns, along side
walks, in a city softball field, a grass parking lot, and a cemetery. All of the
sites are to some degree disturbed and other vegetation consists primarily
of mixed grasses (predominately Cynodon dactylon (L.) Pers., Paspalum notatum
Fluegge, and Eremochloa ophinroides (Munro) Hack.) and other bulb species
([pheton uniflorum (Lindley) Raf., A//inm bivalve (L.) Kuntze, and Allivm canadense
L.). Copper Lily is so abundant there chat in some lawns it forms a com-
plete ground cover when in foliage. The soils are sandy and well drained.
Flowering of Copper Lily is most common in July, August, and September,
when periods of drought are broken by summer thunder showers. The foli-
age appears in late autumn, usually in October, and remains green throughout
the winter. The total area occupied by Copper Lily in Troy is about three to
five acres, and extends in a unbroken population for 276 meters along Three
Notch Street.

In China Grove and in the High Ridge area Copper Lily is found on roadsides
and in lawns. It is associated with many of the same species as the popula-
tion in Troy, with Cynodon dactylon (L.) Pers. and Paspalum notatum Fluegge

Sipa 18(1): 353. 1998

Sipa 18(1)

354

LAUDERDALE MADISON S JACKSON

}
oe

load V7

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RANDOLPH

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Fic. 1. Map of Alabama with locations of Pike and Bullock counties.

predominating. The soils in China Grove and in High Ridge are coarse sands
and the sites tend to be xeric. Undisturbed areas nearby support a dwarf
oak-lichen sandhill community. The population in China Grove extends
for 301 meters along Pike County Highway 37 and the High Ridge popu-
lation extends for 142 meters along Bullock County Highways 7 and 14.
Estimates of total population size are three acres for China Grove and two
acres for High Ridge.

The fourth population is located about eight miles southwest of Troy
near Goshen. It is also found on a roadside and extends for 75 m along Pike
County Highway 25. The soil is sandy and the main associate at this site 1s
Paspalum notatum Fluegge. This is the smallest population, covering approximately
one-half acre.

Monitoring of a planted population of copper lily for five years revealed
that the seed generally fell within 15 cm of the parent plant and that seed-

Sipa 18(1): 354. 1998

Notes 355

lings were readily produced. Seedlings generally flowered for the first time
in their third year of growth without special care.

Holmes and Wells (1980) reinforced the suggestion of H.H. Hume (Sealey
1937) that the distribution of Copper Lily in North America was the result
of human activity. They speculated that the distribution of the Copper Lily
in North America was most likely the result of traveling Spanish mission-
aries as all of the Texas and Louisiana populations are near the sites of Span-
ish missions. As there is no record of a Spanish mission ever having been
located in Troy or in Pike county, it is most likely that the species reached
Alabama by other means. Historical records of the Troy-Pike county area
reveal that in 1865 a man by the name of William Murphree left Troy and
moved to Walker County, Texas. In letters to his mother, Murphree states
that he is enclosing seeds for her to plant. In other records Mrs. Murphree
is noted for planting many of the decorative plants in Troy at that time.
The Murphrees lived on Three Notch Street in Troy, the site of the largest
Copper Lily population reported above. Walker County, Texas, where Mr.
Murphree moved is listed by Holmes and Wells as a place where Copper
Lily was collected (1980). We therefore speculate that Copper Lily was in-
troduced into Alabama by the Murphree family from populations in Walker
County, Texas. As all of the sites reported above are old communities, it is
likely that Mrs. Murphree shared the plant with friends or relatives nearby.

Voucher specimens: ALABAMA: Bullock Co.: T13N R21—22E sect. 36-31, intersec-
tion of Bullock Co. Hwy 7 and 14 and un-numbered dirt road, high ridge, roadsides and
lawns, full sun, deep dry sandy soil, abundant, 12 Jul 1996, A.R. Diamond 10413 (AUA).
Pike Co.: TLON R21E sect. 29, Troy, U.S. Hwy 29 (Three Notch Street), north of down
town at Knox Field, abundant along sidewalks and road, lawns, and vacant lots, full sun,

ay soil, flowered after shower during drought, no leaves seen, 15 Jul 1988, A.R.
Diamond 5210 (AUA); T12N R21E sect. 9, China Grove, just east of Pike Co. Hwy 7,
roadside, full sun, dry sandy soil, common, 17 Jul 1995, A.R. Diamond 9691 (AUA); TIN
R20E sect. 6, Pike County Hwy 25, 0.2 mi S of U.S. Hwy 29, ca. 75 m along the west side
of the road, full sun, flowers yellow, 3 Oct 1997, A.R. Diamond 11131 (AUA).

Duplicates will be distributed at a later date.—A/vin R. Diamond, Jr.,
Charles P. Chapman, and Jim Brummett, Department of Biology, Troy State Uni-
versity, Troy, AL 36082, U.S.A

REFERENCES
BURKHALTER ,J- R. 1984. Additions to the vascular flora of Florida. Castanea 49:181.
HOo.LMEs, d C.J. WeLts. 1980. The distribution of hile ee . Her.)
Traub in ne America and North America- Texas and Lou a. Sida 8:328-333.
SEALY, J.R. 1937. Zephyranthes, Pyrolirion, Habranthus and Hee J. saat Hort. Soc.
62:195—209

Sipa 18(1): 355. 1998

356 Sipa 18(1)
PLANTAGO CORONOPUS (PLANTAGINACEAE) NEW TO TEXAS

Plantago coronopus L., native to Europe and the Mediterranean area (Chater
& Cartier 1976), was reported as naturalized in California (Dempster 1993).
In the eastern United States it has been variously described as rarely adven-
tive on ballast heaps at seaports (Gleason 1952), occurring sporadically about
ports without persisting (Fernald 1950), and as a repeated introduction in
ballast that does not become established (Gleason & Cronquist 1991). It
was not reported in standard floras of the southeastern United States (e.g.,
Small 1913; Radford et al. 1968), or in the floras of Missouri (Steyermark
1963), New Mexico (Martin & Hutchins 1981), Great Plains (McGregor &
Brooks 1986), Arkansas (Smith 1994), Oklahoma (Taylor & Taylor 1994),
or Texas (Correll & Johnston 1970). It was also not included in more recent
works on Texas plants such as Hatch et al. (1990) or Jones et al. (1997), nor
was it listed by Rosatti (1984) in his treatment of Plantaginaceae for the
Generic Flora of the Southeastern United States.

As part of the collecting effort for the forthcoming Shinners & Mahler's
Ulustrated Flora of North Central Texas (Diggs et al., forthcoming), a collec-
tion made in 1998 in Fort Worth (Tarrant County) is apparently the first
documented occurrence of this species for Texas.

Voucher specimen: TEXAS. Tarrant Co.: weedy area in landscape, 301 Crestwood, Fort
Worth, 24 May 1998, O’Kennon 14221 (BRIT).

At the collection locality numerous individuals were observed as landscape
weeds and other individuals were observed in nearby weedy areas. The mode
of introduction is unclear, but seeds were possibly introduced with landscape
plants. Introduction could also have occurred through spread from intentional
cultivation. Since 1997, seeds of Plantago coronopus have been offered by Johnny’s
Selected Seed Company for the specialty salad green market under the name
“Minutina” or “Erba stella” and, until recently, seeds were sold by the Orna-
mental Edibles Seed Company as an “edible landscape plant.” Other north
central Texas populations have not been found and it is unclear whether the
species will become more widely established. However, some individuals were
very robust, with one having 65 inflorescences. Also, the species is self-com-
patible (Delden et al. 1992) and is tolerant to drought, soil compaction, and
competition (Mook et al. 1992). The seeds are highly viable and germinate
well in soils with low moisture content (Blom 1992).

This species, commonly known as buck-horn plantain, differs from all

other plantains occurring in Texas in having deeply pinnately lobed leaves,
short hairs on the corolla tubes, and 3- or 4-locular capsules. Other mem-
bers of the genus occurring in the state have leaves entire or with few teeth
or shallow lobes, corolla tubes glabrous, and 2-locular capsules.

Sipa 18(1): 356. 1998

Notes 357

iy NN ee

=—

——

pepe

we
N

Fic. 1. Habit of Plantago coronopus (with permission from Gleason 1952).

Plantago coronopus (Fig. 1) can be recognized by the following descrip-
tion (Butcher 1961; Gleason 1952; Chater & Cartier 1976; Dempster 1993;
Rahn 1996): Pubescent annual or biennial with a taproot and leaves in a
basal rosette; leaves closely spreading on the ground or ascending, linear to
lanceolate in outline, 4-25 cm long, with spreading-ascending, acute, +
linear lobes, the lobes sometimes toothed or lobed; inflorescences sometimes
numerous, J—50 cm long including the leafless, densely hairy peduncle terminated
by a narrowly cylindric, dense spike to ca. 12.5 cm long; bracts usually not
surpassing the flowers, broadly scarious-margined at base, the keel prolonged
into an acuminate tip; corolla tubes covered with short hairs; petals acute;
stamens 4; capsules 3- or 4-locular; seeds 3—5 per capsule, dimorphic, one
per locule, ca. 1.5 mm long, and one smaller than the others, ca. 0.5 mm
long, occurring in an upper compartment in the ovary on the abaxial side;
flowering late Apr—Sep.

—Robert J. O’Kennon, Botanical Research Institute of Texas, 509 Pecan Street,
Ft. Worth, TX 76102, U.S.A., bokennon@ brit.org; George M. Diggs, Jr., De-
partment of Biology, Austin College, Sherman, Texas 75090, U.S.A. & Botanical
Research Institute of Texas, gdiges@austinc.edu; and Ronald K. Hoggard, Department
of Biology, University of Central Oklahoma, Edmond, OK 73034, rhoggard@ ionet. net.

Sipa 18(1): 357. 1998

358 Sipa 18(1)
REFERENCES

ae om, C.W.P.M. 1992. oo of Plantago populations: Germination and establishment.
PJ.C. Kuiper & M. Bos, eds. Planta A Multidisciplinary Study. Pp. 88-97. Springer-
7 i, Berlin, Heider New Y

Burcuer, R.W. 1 new illustrated eae flora. Leonard Hill Books Limited, Lon-
don, England, a
CorrELL, D.S. and i. Ce Jones 1970. Manual of the vascular plants of Texas. Texas

Research Foundation, Rennet

Carer, A.O. and D. Cartier. 1976, ee In: T.G. Tutin, V.H. Heywood, N.A. Bu urges,
D.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. Flora Europaea 4:38-44

DELDEN, Ww Van, H. VAN Dyk, and K. oo FF. 1992. The genetics of Plantago species. In
P.J.C. Kuiper and M. Bos, eds. — A multidisciplinary study. Pp. 295-309. minets
Verlag, Berlin, Heidelberg, New Yor

Dempster, L.T. 1993. Pastime a J.C.Hickman, ed. The ce manual, higher
plants of California. Pp. 820-821. Univ. of California Press, eley

Diccs, G.M. Jr., B.L. Lipscoms, a R.J. O'KENNON. Shinners & sea illustrated flora
of North Central Texas. To be published in late 1998 by the Botanical Research Insti-
tute of Texas, Fort Worth.

FERNALD, M.L. 1950. Gray's manual of botany, 8th ed. Reprinted 1987. Dioscorides Press,
Portland, OR.

GLeason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United
Scates and adjacent Canada, 3 vols. New York Botanical Garden, B

and C RONQUIST, A. | 991. Manual of the vascular plants . northeastern United

States and adjacent Canada, 2nd ed. Van Nostrand Reinhold Company, New York.

Hare H, S.L., K.N. GANbuI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas.

Qo

N

Texas Agric. Exp. Sta. Misc. Publ. 1655:1-158.
JONES, S. D. oe WaRke siete Monrcomery. 1997. Vascular plants of Texas: A compre-
bibliography, and index. Univ. of Texas Press, Austin.

Martin, W.C. and C.R. Aurcuine. L981. A fier of New Mexico, 2 vols. J. Cramer, Ger-

many

McGrecor, R.L. and R.E. Brooks. 1986. ins oe In: Great Plains Flora Associa-
tion. Flora 7 the Great Plains. Pp. 742-747. Univ. Press of Kansas, Lawrenc

Mook, J.H., J. Harck, J. WAN DER Toorn, and PH. VAN TIENDEREN. 1992. Ecology of Plantago

populations: =e demographic structure of populations. In: P.J.C. Kuiper and M. Bos
eds. Plantago: A multidisciplinary study. Pp. 69-87. Springer-Verlag, Berlin, peel:

berg, New York

Raprorb, A.E., H.E. Anes, and C.R. Bett. 1968. Manual of the vascular flora of the Caro-
linas. The Univ. of North Carolina Press, Chapel Hill.

Rann, K. 1996. A phylogenetic study of the Plantaginaceae. J. Linn. Soc., Bot. 120: 145—
198,

Rosati, oa 1984. The Plantaginaceae in the southeastern United States. J. Arnold Arbor.
65: 533-562.

SMALL, 1K 1913. Flora of the southeastern United States, 2nd ed. Published by the au-
thor, New York.

SmitH, E.B. 1994. Keys to the flora of Arkansas. Univ. of Arkansas Press, Fayetteville.

STEYERMARK, J.A. 1963. Flora of Missouri. The Iowa State Univ. Press, Ames.

Taytor, R.J. and C.E.S. Taytor. 1994. An annotated list of the ferns, fern allies, gymnosperms
and flowering plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Univ., Durant.

Sipa 18 (1): 358. 1998

Novtes 359
PTERIS VITTATA (PTERIDACEAE), A NEW FERN FOR TEXAS

Preris vittata L., commonly known as ladder brake, Chinese brake, or Chi-
nese ladder brake, is an Asian native well known as an escape from cultiva-
tion in coastal areas of the southeastern United States. The species was not
included in Small’s 1903 or 1913 treatments of the southeastern flora. Small
later (1938) treated the species as Pycnodoria vittata (L.) Small, indicating
that it had been collected in Florida many years previously and that during
the 1930s it was abundant at several Florida localities (e.g., Everglades).
He also indicated that it was established in Alabama, Louisiana, the West
Indies, and South America. Currently, Preris vittata is common in southern
Florida where it can be found in almost every habitat with exposed lime-
stone (e.g., pinelands) and on a variety of man-made calcareous substrates
(e.g., sidewalks, buildings, old masonry) (Nauman 1993). Radford et al.
(1968) cited a South Carolina record and noted that it was a rare escape.
Nauman (1993) mapped the species as occurring from southern South Carolina
south to Florida and west to eastern Louisiana; he also showed isolated oc-
currences in southern California and the District of Columbia.

Preris vittata was not included in the Texas flora by Correll (1956, 1966),
Correll and Johnston (1970), Stanford (1976), Hatch et al. (1990), or Jones
et al. (1997), nor has it been reported from Oklahoma (Taylor & Taylor 1994).
It is now known from Texas based on the following collection:

Voucher collection: Texas. San Saba Co.: ca. 3 mi N of Bend, on the Edwards Plateau,
growing from a limestone boulder with Adiantum captllus-veneris L. at edge of Rough Creek
(tributary of the Colorado River), 6 Nov 1987, ].W. Stanford 5308 (BRIT, HPC, SPLT).

Subsequent flooding has destroyed the site. The identification was con-
firmed by Jim Blassingame (South Plains College, Levelland, TX).

Of the five Pteris species in the United States (most occurrences are in
the southeastern part of the country), four are introduced and one, P. bahamensis
(J. Agardh) Fée, is native to south Florida (Nauman 1993). Besides P. vittata,
only P. multifida Poir. (spider brake, Chinese brake, Huguenot fern), is known
from Texas. This widely cultivated native of China is naturalized in east
and southeast Texas and the Rio Grande Valley (Correll & Johnston 1970;
Hatch et al. 1990). The deeply palmately 3-divided pinnae (at least some)
of P. multifida are quite distinctive.

Preris vittata can be recognized and distinguished as follows: stems short-
creeping, stout, densely scaly; leaves clustered, strictly 1-pinnate (the pin-
nae without lobes or divisions), 0.3—1 m long including petiole; pinnae 12—
20(—30) pairs per leaf, linear-lanceolate to linear-attenuate, 2-18 cm long,
4—9 mm wide, long-attenuate or sharply acute apically. ically cordate
to widened or truncate basally; serrulate matginally: distal pinnae conspicuously

Stipa 18(1): 359. 1998

360 Sipa 18(1)

longer than proximal pinnae (leaf blades thus oblanceolate in outline) with
the terminal pinna typically longest; petioles and often rachises densely scaly;
sori usually of a continuous narrow band near the margins of the abaxial
surface of the pinnae (thus most of the abaxial blade surface exposed) (Long
& Lakela 1971; Nauman 1993; Smith & Lemieux 1993), The species is a
tetraploid, with 27 = 116 (Nauman 1993). An illustration and a detailed
description are available in Small (1938). A recent treatment of Preris, in-
cluding a key to taxa occurring in the he States and an illustration of
P. vittata, can be found in Nauman (1993

—Jack W. Stanford, Department of Biology, H. a Payne University, Brownwood,
TX 76801, U.S.A., jstanfor@hputx.edu and George M. Diggs, Jr, Department
of Biology, Austin College, Sherman, TX 75090, U.S.A. & Botanical Research
Institute of Texas, 509 Pecan Street, Ft. Worth, TX 76102, U.S.A., gdiggs@austine. edu,

REFERENCES

Correétt, D.S. 1956. Ferns and fern allies of Texas. Texas Research Foundation, Renner.

1966. sale In: C.L. Lundell, ed. Flora of Texas 1:3—121. Texas
‘Research Foundation, Renn

and M C Jonsson 1970. Manual of the vascular plants of Texas. Texas

Research Poundaciun Ren

Hatcu, S.L., K.N. GANDHI, a L E. Brown. mee Checklist of the vascular plants of Texas.
Texas Agric. Exp. Sea. Misc. Publ. 1655:1 :

Jones, §.D., J.K. Wiper, and P.M. NoMeakee 997. Vascular plants of Texas: A comprehen-
sive checklist including synonymy, bibliography, and index. Univ. of Texas Press, Austin.

Lona, R.W. and O. Lakers. 1971. A flora of tropical Florida. Univ. of Miami Press, Coral

a

a ,F
Nauman, C.E. 1993. Preris. In: Flora of North America Editorial Committee, eds. Fl. North
Amer, 2:132-—135. Oxford Univ. Press, New York and Oxforc
Raprorp, A.E., H.E. Anzes, and C.R. Bri, 1968, Manual of the oasculat flora of the Caro-
linas. The Univ. of North Carolina Press, Chapel Hill.
SMALL, i K. 1903. Flora of the southeastern United States. Published by the author, New
Yor

. 1913. Flora of the southeastern United States, 2nd ed. Published by the
author, New York.
1938. Ferns of the southeastern United States. The Science Press Printing

Co., Lancaster, PA.

SmitH, A.R. and T. Lemipux. 1993. Preridaceae. In: J.C. Hickman, ed. The Jepson manual,
higher plants of California. Pp. 439-444. Univ. of California Press, Berkele

STANFORD, J.W. 1976. Keys to the vascular plants of the Texas Edwards Plateau and ad ja-
cent areas. ee by the author, Brownwood, TX

Tavior, R.J. and C.E.S. Taytor. 1994. An annotated tis of the ferns, fern allies, gymno-
sperms and Tower plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Uni-
versity, Dur.

Sipa 18(1): 360. 1998

Notes 361

LYCIANTHES ASARIFOLIA (SOLANACEAE)
NEW AND WEEDY IN TEXAS

In July of 1997 an unknown plant was collected in Houston and sent to
Texas A&M for identification. It proved to be Lycianthes asarifolia (Kunth
& Bouché) Bitter, a member of the Solanaceae native to South America. This
species has been previously known in North America only from New Or-
leans, Louisiana (Darwin & Feibelman 1991). De Rojas and D’Arcy (1997)
incorrectly cited that collection as being from Texas.

This species is recognizable by its prostrate habit, having slender stolons
that root at every node. The leaves are strongly cordate, entire, and long-
petioled. Sometimes a second leaf is produced at non-flowering nodes: if
the primary leaf is relatively large, the second leaf is usually small and of-
ten stipule-like; when the primary leaf is relatively small, the second is of-
ten nearly the same size. The flowers are white, rotate, and ca. 1.5 cm broad.
The yellow anthers are connivent around the style and poricidally dehis-
cent. The fruit is an edible reddish-orange berry ca. 1.3 cm in diameter,
closely subtended by the truncate calyx. Plants are self-infertile (Dean 1997).
A good illustration can be found in the article by De Rojas and D’Arcy and
images of the Houston plants can be seen at Texas A&M’s Bioinformatics
Working Group Image Gallery page (http://www.csdl.tamu.edu/FLORA/
imaxxsol.htm). Note that the leaves of the Houston plants are nearly all
rounded apically rather than pointed as in the article illustration.

A visit to the collection site revealed that this plant has overrun several
residential yards in Houston, forming a dense, attractive ground cover in
shaded areas. It is apparently reproducing both vegetatively and sexually,
for numerous flowers and immature fruits were seen in early December 1997
and mature fruits were collected in March 1998. (Darwin and Feibelman
found no fruit in the Louisiana population and none has since been seen on
plants grown from cuttings taken from that population.) The authors also
suspect that propagules are being carried from yard to yard on the equip-
ment of landscape maintenance companies. The exact time and point of
introduction is not known, but the population has been in existence for three
or more years. Residents’ attempts to control the plant by hand-pulling,
mowing, herbicide application, and removal of infested sod have proved
unsuccessful. It appears to be tolerant of Houston’s winter weather and suffers
only partial dieback during the hottest summer days. Should this plant become
established in nearby Memorial Park (a large, wooded area), it could be nearly
impossible to eradicate. Herbicide trials were begun in one of the Houston
yards in March of 1998. After two rounds of tests, no treatment has pro-

Siva 18(1): 361. 1998

362 Stipa 18(1)

vided complete control, and treatments which have weakened the Lycianthes
have also damaged the surrounding turf.

It is possible that this plant could become a noxious weed in southern
Texas. The USDA Animal and Plant Health Inspection Service is conduct-
ing a formal risk assessment. (The Louisiana Department of Agriculture
has considered surveying for the plant in the New Orleans area.) The au-
thors encourage identification, documentation, and destruction of any new
colonies subsequently discovered. Should other populations of this plant
be found, please contact one of the authors.

Voucher specimen: U.S.A. TEXAS. Harris Co.: Houston, shaded residential yard of
502 Lindenwood, with Quercus virginiana, Pinus, Stenotaphrum secundatum, and ning a
in moist sandy loam, 6 Dec ee Mary Ketchersid 1 20697- A (BH, BRIT, MO, NO, NY.
TAES, TAMU, TEX, US, WAVI).

We thank Michael Nee (NY) for his assistance in identifying this plant.
—Monique Dubrule Reed, Biology Department, Texas AGM University, College
Station, TX 77843-3258, U.S.A. and Mary Ketchersid, Agricultural and En-
vironmental Safety, Texas Agricultural Extension Service, College Station, TX 77843-
2488

REFERENCES
Darwin, S.P and T. FeIBeLMAN. 1991. Lycianthes asarifolia (Solanaceae) new to North America.
Sida 14:605—606.
Dean, E. 1997. Personal communication. Dr. Dean studies the taxonomy of Lycranthes at
the aa of California- Davi
De Rojas, C.E. and W. 1997. The genus Lycianthes (Solanaceae) in Venezuela.
rete pee Bot. Gard. 84 se —200.

5
joe

Stipa 18(1); 362. 1998

NOoTrESs 363

FIRST REPORT OF THE GENUS BURMEISTERA
(CAMPANULACEAE) FROM HONDURAS

Burmeistera Triana (Campanulaceae: Lobelioideae) is a genus of 96 species
endemic to the Neotropics (Lammers 1998); it was last monographed by
Wimmer (1943). The genus is closely related to two other Neotropical endemics,
Centropogon C. Presl and Siphocampylus Pohl (Pepper et al. 1997; Lammers
1998). All are large robust plants (herbaceous or suffruticose perennials,
shrubs, subshrubs, or lianas) with large flowers (corollas averaging 30—60
mm long or more) borne singly in the axils of the upper leaves or aggre-
gated into terminal bracteate or foliose (sometimes corymbiform) racemes;
the tube of the corolla is neither fenestrate nor cleft dorsally and if the lobes
are dimorphic, it is the two dorsal ones that are the larger. Burmezstera 1s
distinguished from its allies by its combination of usually ebracteolate pedicels,
green or yellow corolla often suffused with maroon or purple, large falcate
or reflexed dorsal corolla lobes, the wide open orifice of the anther tube,
baccate often inflated fruit, and oblong to fusiform seeds much longer than
road.

Most of the species of Burmeistera are found in montane areas from Costa
Rica to Ecuador. At the southern limits of this range, two Ecuadorean spe-
cies extend south into Peru (Stein 1987). At the northern end, a single spe-
cies is known to occur north of Costa Rica: Burmerstera virescens (Benth.) Benth.
& Hook. ex Hemsl. It has been reported (McVaugh 1943; Wimmer 1943;
Nash 1976) only from Guatemala, where it occurs in the departments of
Alta Verapaz, Baja Verapaz, Huehuetenango, Quezaltenango, San Marcos,
and Suchitepéquez. This is a disjunction from its congeners of nearly 700
km. That gap has been narrowed, however, by the discovery of this same
species in central Honduras. This is the first report of any member of Barmezstera

from that nation.

Voucher specimen: HONDURAS. Depto. Olancho: Parque Nacional la Muralla, trail
above Visitors’ Center, ca. 14 km above La Union, 15°05'N 86°40'W, in dense high pri-
mary Quercus forest, 1400 m, 27 Oct 1996, PJ.M. Maas & H. Maas 8441 (U).

The new locality in Honduras is approximately 350 km east of the near-
est conspecific populations in Guatemala. The Qvercus-dominated primary
forest there was quite rich in epiphytes, including Plewrothallis tuerckheimii
Schlecht. (Orchidaceae), Columnea rubrecaulis Stand|. (Gesneriaceae), and Peperomia
hoffmannii C. DC. (Piperaceae). Understory shrubs included Besleria solanoides
Kunth (Gesneriaceae), Monnina ferreyrae Taylor (Polygalaceae), Tournefortia
sp. (Boraginaceae), scandent Sphyrospermum majus Griseb. (Ericaceae), and a
treelet species of C/ethra (Clethraceae). Species found in the herbaceous layer

Sipa 18(1): 363. 1998

364 Stipa 18(1)

were Psilochilus macrophyllus (Lind\.) Ames (Orchidaceae), Renealmia mexicana
Klotzsch ex Peterson (Zingiberaceae), Smilacina sp. (Asparagaceae), Spige-
lia sp. (Spigeliaceae), the saprophyte Gymmnosiphon suaveolens (Karst.) Urb.
(Burmanniaceae), and the root-parasite Monotropa uniflora L. (Monotropaceae).
Adjacent patches of secondary forest contained small trees of Sawrania veraguensis
Seem. (Actinidiaceae), shrubs of Trivmfetta speciosa Seem. (Tiliaceae), and the
herbs Canna tuerckheimii Kraenzl. (Cannaceae) and Hydrocotyle mexicana Cham.
& Schlecht. (Apiaceae).

—Thomas G. Lammers, Department of Botany, Field Museum of Natural H1s-
tory, Chicago, IL 60605-2496, U.S.A.; and P.J.M. Maas, Department of Plant
Ecology and Evolutionary Biology, Herbarium Division, Willem C. van Unnikgebouw,
Heidelberglaan 2, 3584 CS Utrecht, Netherlands.

REFERENCES

Lammers, T.G. 1998. Review of the Neotropical endemics Burmeistera, Centropogon, and
Siphocampylus (Campanulaceae: Lobelioideae), with description of eighteen new specie
and a new section. Brittonia 50:233—262.

McVauGu, R. 1943. Campanulaceae (Lobelioideae). N. Amer. Fl. 32A:1—

Nasu, D.L. 1976. Campanulaceae. In: Flora of Guatemala, part EX, no. 4. a Mus. Nat.

, Bor. ae a 276-431.

PEPPER, ae H.G. Gustarsson, and V.A. ALBERT. 1997. Molecular systematics of
Nepal earn (Campanulaceae), with emphasis on Burmeistera, Centropogon,
anc ene us, and S utility of oe and floral characters in lobelioid classification
{abstract}. Amer. J. Bot. 84(6, Suppl

STEIN, B. 2 1987. seater ‘ofthe genus cee (Campanulaceae: Lobelioideae) in Peru.
Ann. Missouri Gard. 74:494-496.

se FE. 1943. Campanulaceae-Lobelioideae, I. Teil. In: R. Mansfeld, ed. Das Pflanzenreich,

WAN Ts) oF Verlag ¥ isa Engelmann, Leipzig

SipA 18(1): 364. 1998

Nores 365

TRIRAPHIS MOLLIS (POACEAE: ARUNDINEAE) A SPECIES
REPORTED NEW TO THE UNITED STATES

Prior to this report the occurrence of Triraphis and T. mollis R. Br. (purple
plumegrass) in naturalized populations was not documented in the United
States (Chase 1951; Correll & Johnston 1970; Gould 1975; Hatch et al.
1990; Kartesz 1994; Jones et al. 1997). Specimens of this taxon were first
collected in naturalized populations by William Godwin on 15 Mar 1993
in Dimmit County. Since the original U.S. collection, William E. Fox HI
collected the species in the same area on | May 1996.

Cultivated specimens collected from an experiment station in Biloxi,
Mississippi were located in TAES. However, specimens have not been lo-
cated that were from escaped or naturalized populations. TROPICOS has
reported this species as being present in the U.S. because of a listing by the
U.S. Department of Agriculture in their “National List of Scientific Plant
Names” (U.S.D.A. 1982). In that U.S.D.A. publication, the distribution
was not listed because the plant species was not naturalized.

The species was identified to genus using Clayton and Renvoize (1986)
and to species using Simon (1993). The identification was then verified using
the Tracy Herbarium (TAES) plant specimens from Australia and specimens
from U.S. National Herbarium (US). Lazarides (1970) reported the grass
to be grazed in Australia, but stated that it appeared to be less valuable
than many short-lived perennials. Clayton and Renvoize (1986) reported
the genus to be allied with Neyraudia because of gross morphology and the
slender microhairs.

Superficially, 7’ mollzs resembles Aristida. However, upon examination
of the spikelets with a dissecting microscope the following observations become
apparent. Spikelets have several florets with the reduced floret apical. Also
the lemmas are 3-veined, 3-awned. The lemma awns are extensions of the
3-veins of the lemmas. In T: mo//is, the central lemma awn originates from
the lemma apex and lateral awns below and on both sides of the central
awn, whereas in Aristida the lemma awns originate from a column or the
lemma apex and branch from a common point of origin.

Triraphis mollis R.Br., Prodr. Fl. Nov. Holl. 185. 1810. (Fig. 1).

Caespitose perennial. Culms to 50 cm tall, erect to ascending, unbranched;
nodes glabrous, purplish black; internodes solid. Leaves mostly basal; sheaths
glabrous, rounded, margins free; auricles minute; ligules a fringe of hairs
about | mm long; blades 6-17 cm long, 1-2 mm wide, flat becoming in-
volute, glabrous. Panicles 7-20 cm long, 8-15 mm wide (excluding awns),
contracted, somewhat interrupted, purplish tinged. Spikelets 4-12 mm long,

Stipa 18(1); 365. 1998

366

Sipa 18(1)

\ Mo Hy;
\ i] WY

RAY

Fic. 1. Triraphis mollis. A. habit; B. spikelet, glumes separated from florets; C. floret, palea

view with lemma margin and the associated ae aged D. ligule.

Stipa 18(1): 366. 1998

Nores 367

crowded, pedicellate, laterally compressed, disarticulation above glumes and
between florets; florets 4-10, callus bearded, reduced florets 1—2 and api-
cal. Glumes subequal (not consistent in which glume is longer), 1-veined,
hyaline, shorter than spikelets, awnless; first glumes 3.5—5 mm long; sec-
ond glumes 4—5.5 mm long; lowermost lemmas 4—5 mm long, 3-veined
and 3-awned, deeply bifid with teeth to | mm long, membranous, median
vein awns 6-9 mm long from between teeth of bifid apex; lateral vein awns
5.5-7 mm long, divergent; lateral veins with long pilose hairs (hairs occa-
sionally on the lemma body); lowermost paleas 2—2.5 mm long, 2-keeled,
hyaline, glabrous to scaberulous. Stamens 3; anthers 0.2—0.3 mm long, yellow.

Collection localities were characterized by deep sandy soils of the Brystal
Fine Sandy Loam (Stevens & Arriaga 1985). Historically, well drained sandy
soils derived from Carrizo Sandstone were called Norfolk Fine Sand (Carter
1931). These soils are derived from parent material in the Carrizo Sands
Formation. Carrizo Sand has been shown to have hygroscopic properties that
ameliorate the effects of a xeric climate and consequently influence its plant
formation (McBryde 1933).

Specimens examined: AUSTRALIA. New South Wales: Sydney; 18??, J. Maiden s.n.
(TAES). Northern Territory: Simpson Desert, ca 15 km N of Andado Homestead (250
km SE of Alice Springs), 11 Jul 1968 J. Weber 958 (TAES). Queensland-Warrego Dis-
trict: Charleville, cemetery, Oct-Nov 1945, M. Clemens s.n. (TAES).

UNITED STATES. Texas-Dimmit Co.: 14 mi NW of Carrizo Springs on Cometa Road,
near sandy dirt road between Cometa and ral 277,15 Mar aes W. Godwin s.n. a
13 mi W of Carrizo Springs, 2 mi W of FM 393 along Hwy 277,

Road, along W side of road, 1 May 1996 WE. Fox III, 548A (TAES). eee with rr eae

Pennisetum, Setaria, Acacia and Prosopis.

ACKNOWLEDGMENTS

Financial support was provided in part by Texas Higher Education Co-
ordinating Board—Advanced Research Program and the Texas Agricultural
Experiment Station.

—Stephan L. Hatch, William E. Fox Ul, and John E. Dawson LI, 8.M. Tracy
Herbarium, Department of Rangeland Ecology and Management, Texas AGM University,
College Station, TX 77843-2126 U.S.A.; and William B. Godwin, Department
of Entomology, Texas AGM University, College Station, TX 77843-2475 U.S.A.

REFERENCES
Carter, W.T. 1931. The soils of Texas. Texas Agric. Exp. Sta. Bull. 431.
Cuase, A. 1951. Hitchcock’s manual of the grasses of the United States, 2nd. ed., U.S.D.A.
Misc. Publ. No. 200. U.S. Government Printing Office, Washington, D.C

Stipa 18(1): 367. 1998

368 Sipa 18(1)

Cayton, W.D. and S.A. REeNvoize. 1986. Genera graminum, grasses of the world. Kew
Bull. Addit. ce XIII. Her Majesty's Stationary Office, London.

Correit, D.S. and M.C. JOHNSTON.
Resaten Foundation, Renner.

GouLb, . W. 1975. The grasses of Texas. Texas A&M University Press, College Station.

Harcn, $.L., K.N. GANDHI, and . = Brown. 1990. Checklist of the vascular plants of Texas.
Texas oe Exp. Sta. Bull. MP-1655.

Jones, $.D., J.K. ca ue 1997. Vascular plants of Texas, a compre-
hensive checklist including synonymy, bibliography and index. University of Texas Press,
Austin,

Karrtesz, J.

1970. Manual of the vascular plants of Texas. Texas

1994, A synonymized checklist of the vascular flora of the United States, Canada
and Greenland. scciah of North Carolina Press, Chapel Hill.

Lazaripes, M. 1970. The grasses of central Australia. Australian Natl. Univ. Press, Canberra.

McBrype, J.B. 1933. The vegetation and habitat factors of the Carrizo Sands.
Monogr. 3:247-297

Simon, B.K. 1993. A key to Australian grasses. Queensland Department of Primary In-
dustries, Brisbane, Australia

STEVENS, J.W. and D, ARRIAGA. 1985. Soil survey of Dimmit and Zavala counties, Texas.
USDA S.C.S and Texas Agricultural Experiment Station.

Ecological

Sipa 18(1): 368. 1998

PRIORITY OF THE NAME AGALINIS HARPERI
(SCROPHULARIACEAE) OVER THE NAMES AGALINIS
DELICATULA AND AGALINIS PINETORUM

This paper is part of an ongoing study by the author of the vascular plant
genus Aga/inis along the Gulf Coastal Plain. The objective of this note is to
clarify one of the many nomenclatural difficulties associated with North
American species of this genus.

Pennell (1913) described three species of Aga/inis from Florida: A. delicatula,
A. harperi, and A. pinetorum. These species have recently been shown to be
conspecific (Canne-Hilliker and Kampny 1991). Later, in 1929, Pennell
reduced A. delicatula to a variety of A. pinetorum as A. pinetorum vat. delicatula
(Pennell) Pennell, considering it to be a weak variety at best and stating in
his description that it was “Of doubtful rank” (Pennell 1929). In their study,
Canne-Hilliker and Kampny (1991) concluded that that all three names
represented one species and and chose A. harper for this complex. Kartesz
(1994), although apparently aware of the work of Canne-Hilliker and Kampny,
chose instead to combine the two species under the name A. pinetorum.
Although Pennell later recognized the name Aga/inis pinetorum vat. delicatula
(Pennell) Pennell at the varietal rank, it must still be taken into account for
nomenclatural purposes because the name A. de/icatula has equal priority
with A. pinetorum (Pennell 1913).

The name Agalinis harperi and a brief description of the species first ap-
peared in the Flora of Miami, published in 1913 (Small 1913), in which
Pennell prepared the treatment of Aga/inis. On 13 August 1913, the August
issue of Bulletin of the Torrey Botanical Club was published, and in this issue
Pennell published for the first time the description of A. delicatula and A.
pinetorum (Pennell 1913). It is clear that Pennell realized that Small’s flora
had appeared (or would appear) before the August 1913 publication of the
Bulletin, this is because he (p. 426) noted that A. harper? was first described
in the Flora of Miamz. But due to the fact that Small published the Flora of
Miami himself, I have been unable to determine an exact publication date
for it. However, in accordance with Art. 31.1 of the Code (Greuter et al.
1994), A. harperi was effectively published prior to publication of the 13
August 1913 issue of Budletin of the Torrey Botanical Club. The valid publi-
cation date is fulfilled in two ways in accordance with Articles 29.1 and
31.1 of the Code:

1. The preface of Small’s Flora of Miami is dated 26 April 1913 (Small
1913); this is the only exact date found in the flora. According to Art. 31.1
of the Code, in the absence of any other evidence validating the date of
a work, the earliest date indicating publication is to be accepted as the

Stipa 18(1): 369. 1998

370 Sipa 18(1)

correct publication date.
1997 communication with John F. Reed, Director of the LuEsther

T. Mertz Library at the New York Botanical Garden, I was able to verify
that the publication of Small’s Flora of Miami did in fact precede the 13
August 1913 publication date of Bulletin of the Torrey Botanical Club—a copy
of Small’s Flora of Miami at the New York Botanical Garden’s library is
date-stamped 21 June 1913. Thus, publication of Aga/inis harperi in Small’s
Flora of Miami obviously preceded the 13 August 1913 publication of
A. delicatula and A. pinetorum in the Bulletin. The valid name for this
species and its synonyms is as follows:
Agalinis harperi Pennell, in Small, Flora of en L762 1913.

Agatinis delicatula Pennell, Bull. kee Bot. Club 40:425. 1913. Agalinis pinetorum var.

delicatula (Pennell) Pennell, >, Acad. Nat. Sci. see) shia 81:172-173. 1929.
Goan var. eee nnell) Pennell, Acad. Nat. Sci. Philadelphia Monogr.

A, neon ae Bull. Torrey Bot. Club 40: ae 25. 1913.
Gerardia harperi (Pennell) Pennell, Acad. Nat. Sci. ladelphia Monogr. 1:441. 1935.
G. piulchella (Pennell) Pennell, Acad. Nat. Sci. ae phia Monogr. 1:440. 1935.

ACKNOWLEDGMENTS

I thank John Thieret of Northern Kentucky University for reviewing the
manuscript and John F. Reed of the LuEsther T. Mertz Library at the New
York Botanical Garden for his assistance. I also thank Judith Canne-Hilliker
of the University of Guelph in Ontario, Canada for sharing her unsurpassed
knowledge of Agalinis with me.—John F. Hays, Ozark Environmental Con-
sulting, 3308 Keenes Edge Drive, Columbia, MO 65201, U.S.A

REFERENCES

CANNE-HIUILLIKER, J.M. and C.M. Kampny. 1991. Taxonomic significance of leaf and stem
anatomy of Aga/inis (Scrophulariaceae) from the U.S.A. and Canada. Canad. J. Bot. 69:1935-
L950,

GreEUTER, W., ER. Burpet, W.G. CHALONER, V. DEMOULIN, D.L. Hawkswortu, P.M. JORGENSEN,
D.H. Nicotson, P.C. Litva, P. TREHANE, and J. McNett. 1994. International code of bo-
ene nomenclature (Tokyo Code) adopted by the fee — Botanical Congress,

<ohama, August-September 1993. Regnum Veg

Pre J.T. 1994. A synonymized checklist of ie ede ar flora of the United States,
Canada, and Greenland, 2d ed. Timber Press, Portland. 2 vols

PENNELL, F.W. 1913. Studies in the Agalinanae, a subtribe of the Rhinanthaceae II. Species
of the Atlantic coastal plain. Bull. Torrey Bot. Club 40:401-439.

PENNELL, F.W. 1929. Aga/inis and allies in North America-IJ. Proc. Acad. Natl. Sci. Phila-
delphia 81:11-249.

PENNELL, F.W. 1935. The Scrophulariaceae of eastern temperate North America. Acad. Nat.
Sci. Philadelphia Monogr. 1:419-476.

SMALL, J.K. 1913. Flora of Miami. Published by the author, New York.

Sipa 18(1): 370. 1998

HEINZ DIETRICH LUBRECHT (1908-1997),
BOTANICAL BOOKSELLER AND FRIEND

WILLIAM R. BURK
Biology Library
University of North Carolina
Box 3280 Coker Hall
Chapel Hill, NC 27599-3280 U.S.A.

Heinz (Harry) Dietrich Lubrecht, age 88, well known botanical and natural
history bookseller, antiquarian book expert, appraiser, and former execu-
tive of the publishing company, Stechert-Hafner, died 12 November 1997,
at the Columbia Presbyterian Hospital, New York City, after suffering a
stroke. Harry, as he was affectionately known by friends, colleagues, and
family, devoted nearly 70 years to the scholarly book trade, particularly in
botany and natural history. In 1974 he established Lubrecht & Cramer, Booksellers
and Publishers, currently located in Port Jervis, NY. His high standards of
business, steadfast honesty, and breadth of knowledge of botanical litera-
ture earned him national and international respect as a bookman. With the
passing of Harry Lubrecht the botanical and bookselling community has
lost a faithful friend and serious scholar.

Born 2 December 1908, in Reutlinger, Germany, Harry was the son of
Adolph August Karl Friederich Wilhelm and Sophie Amalie (Grueninger)
Lubrecht. His father, an architect, was the son of Lutheran Pastor Adolph
Lubrecht and Emma Karoline Kuebel. His mother, well-educated and adept
in three foreign languages, was the daughter of Johann Matthaus Grueninger,
Professor of Mathematics in the Reutlinger Hochschule, and Sophie Kaemerer.
When Harry was five, his parents separated. Subsequently, Harry, his mother,
and two sisters moved to Stuttgart to live in an aunt’s apartment home. He
attended Friedrich Eugens Oberrealschule and completed Hochschule in
Stuttgart. Throughout his school years Harry was an avid reader. In order
to ensure enrollment at his school, Harry secured an annual scholarship by
placing in the top five of his class.

Upon graduation at age 17, Harry became an apprentice in Brueninger’s
department store as part of his continued schooling. His flare for business

was well recognized by the store’s management and in three years he earned
the salary of a regular employee. Refusing to join the Nazi party, he was
compelled to leave Germany. Harry emigrated to the United States where
his Uncle Christian Grueninger, the mayor of Valhalla, NY, would sponsor
him. Arriving in America on 15 May 1928, he soon found a job as a grave

Sipa 18(1): 371. 1998

472 Sipa 18(1)

digger at the local cemetery in Mt. Kensico, NY. When this seasonal work
concluded that fall, Harry looked for employment in New York City. In
October 1928, G.E. Stechert & Co.,! well-known international book firm,
hired him at a weekly salary of $14.00. The stage was thus set for a distin-
guished, life-long career in the book world, a profession practiced by an-
cestors on two sides of his family. Some of the Kuebels were medieval scribes,
the book “publishers” of their time. In the early nineteenth century two
Lubrecht brothers, Heinrich and August Adolph, separately developed ca-
reers as booksellers in Heilbronn and Blaubeuren, Germany. In the mid-
nineteenth century August's son, Karl Theobold, emigrated from Blaubeuren
to Brooklyn, NY. Here he Anglicized his name to Charles T. and founded
with his cousin the publishing company Haasis and Lubrechte.

At Stechert Harry advanced from his initial position as billing clerk to
assistant manager of the Antiquarian Department (April 1936). He was a
natural for this job as he was familiar with German, French, Latin, and Greek.
His facility for languages assisted him in compiling book catalogs and un-
derstanding the contents of many scientific books, especially those in Ger-
man, acquired by the company. Since his immediate boss lacked signifi-
cant experience in the antiquarian book market, Harry enjoyed the freedom
and breadth to develop the antiquarian department about which topic he
read widely. Under his supervision the process of searching for and supply-
ing out-of-print books was significantly accelerated (Anonymous 1952). Among
Harry's promotions were to: Chief of Reference and Customer Service, G.E.
Stechert & Co. (spring 1945); Division Chief, Antiquarian and Out-of-Print
Department, Stechert-Hafner, Inc. (fall 1951); Vice-President and Editor,
Hafner Publishing Co. January 1955); and Vice President, Macmillan Publishing
Co.-Hafner Publishing Division (1969).

After conducting a four-year study on subject publishing, Harry con-
cluded that few firms published books on botany. Through Harry’s efforts
a new subsidiary called Stechert-Hafner Service Agency was established in
1960. Botanists and natural historians are fortunate to have had Harry's

'A brief chronology of this book firm provides an historical ay tive. Stechert & Wolff,
wee on - September 1872, was renamed Gustav E. Stechert (1876), G.E. Stechert (1897),
and G echert & Co. (1904). On | October 1946, ae GE Stechert partnership was
caer a Stechert-Hafner, Inc. (SH) was created with its publishing and reprinting
activities transferred to the newly established Hafner ap Co. In 1960 Stechert-

Hafner Service Agency was established as a subsidiary of SH. Crowell Collier Macmillan,
Inc. (now Macmillan, Inc.) acquired SH and its Cie (1969). Beginning January
1974, the corporate name, Ste ch Vert Macmillan, I Inc., became effecti LIVE: , 1980 Macmillan,

Inc. was divided into three parts of which Macmillan Publishing Co., Inc. contained the
division now called Hafner Press

Sipa 18(1): 372. 1998

Burk, Harry Lubrecht 575,

Harry Lubrecht making a presentation at the Botanical Society of America Meeting, His-
torical Section, held at The Ohio State University, Columbus, 11 August 1987. Photo Coney
Ronald L. Stuckey.

wisdom in developing this subsidiary. This new division served scholarly,
scientific and botanical societies by selling their publications and distrib-
uting scientific and botanical books for European publishers on a large scale.
The Agency distributed publications for the New York Botanical Garden,
Missouri Botanical Garden, Texas Research Foundation, and the Hunt Bo-
tanical Library (Anonymous 1972). During the 1960s and 1970s Harry also
focussed attention on reprinting book classics in botany within the Hafner
Publishing Co. Among the reprint series issued were the Classica Botanica
Americana (CBA) titles by such eighteenth and nineteenth century lumi-
naries as Humphry Marshall, John Torrey & Asa Gray, C. S. Rafinesque,
Stephen Elliott, Thomas Nuttall, and William Darlington. Joseph Ewan,
historian of botany, proposed this series and wrote a bibliographical com-
mentary for each of the reprinted books (Ewan 1967). Ironically, these fac-
simile reprints, as well as many others produced by Hafner, have become as
rare as the originals themselves. The rarity of these volumes was due in part

Sipa 18 (1): 373. 1998

374 Sipa 18(1)

to their limited printing, high demand by scholars, and the destruction of
remaining stock when Crowell Collier Macmillan, Inc. purchased Stechert-
Hafner and its subsidiaries in May 1969.

After this corporate change Harry chose to direct the Publishing Divi-
sion. He could not, however, abandon his interest in and passion for rare
and out-of-print books. Using a substantial part of his lunch hour, he would
work in the Antiquarian Department at noon time. Through his voluntary
work, a veritable labor of love, Harry raised impressive revenues for the firm.

While at Stechert, Harry especially enjoyed appraising private collec-
tions for purchase. He was instrumental in acquiring t ling mathematics

library in 1937, besides many other collections. Representing is company,
Harry participated in the First Antiquarian Book Fair in the United States,

sponsored by the Antiquarian Booksellers’ Association of America and held
in New York City (4-9 April 1960). Confident that the fair would be suc-
cessful, he prepared an attractive catalog (#277) of rare books, the first ever
issued from the Stechert-Hafner Antiquarian Department (Anonymous 1960).

ne of his career highlights was the discovery of 11 first editions of the
works of Linnaeus in a Paris bookshop. These books had an additional his-
torical value since they were eerie by the French botanist and explorer
Michel Adanson (1727—1806), noted for his work on the natural classifica-
tion of plants (Lubrecht 1986).

Harry compiled numerous book catalogs covering general topics, Greek
and Latin authors, and natural history. Although no longer in print, his
catalog (#353), “Early American Botanical Works witha Miscellany of Other
Botanical Rarities” (Lubrecht 1967), provides an historical survey of early
classics in North American botany. This booklet, the result of five years of
searching (Anonymous 1967), contains an introduction (pp. 3-5) by Harry,
an annotated list of 217 botanical titles offered for sale, and a section (pp.
55-60), “Reference Tools for Botany,” by Joseph Ewan. The Missouri Bo-
tanical Garden plans to issue a facsimile reprint of this item as a keepsake
of the XVI International Botanical Congress to be held at the Garden in Sc.
Louis (August 1999), Through the enthusiastic support of Harry, Hafner
Publishing Co. published and assisted in the production of “A Short His-
tory of Botany in the United States,” edited by Joseph Ewan (1969). This
book was distributed at the XI International Botanical Congress in Seattle,
WA, August/September 1969.

Harry retired from Stechert Macmillan on 30 November 1973 and then
briefly compiled catalogs for Jack N. Bartfield, an antiquarian bookseller
of New York City. In December 1974 Harry founded his own business, Lubrecht
& Cramer, Booksellers and Publishers, which was later incorporated (Au-

Sipa 18(1): 374. 1998

Burk, Harry Lubrecht 375

gust 1983). Joerg Cramer (deceased 4 June 1985), a German publisher, specialized
in reprinting classical botanical books, many of which Hafner had distrib-
uted in the United States. Harry’s wife Anne, and later his son Charles, assisted
in the business, originally carried out from their country home in Forestburgh,
NY, and dealing predominantly with new, scholarly books, mostly in botany
and natural history. The company is also a distributor for Koeltz Scientific
Books, Gustav Fischer Verlag, and J. Cramer. Although his bookselling company
primarily was concerned with new titles, Harry continued his profound interests
in rare books and the appraisal of libraries. Among the botanical collec-
tions that he appraised were those of Carroll W. Dodge, the New York
Horticultural Society, Emanuel D. and Ann W. Rudolph, William C. Steere,
and R. Gordon Wasson. The remarkable Rudolph Library of 53,000 vol-
umes of botany and natural history books has since been donated to The
Ohio State University Libraries (Stuessy et al. 1997).

For more than two decades Harry and Anne Lubrecht featured booksale
tables at botanical and scientific meetings and regional mycological forays.
Their service was known not only for the sale of books, typically sold at a
special discount for meeting and foray attendees, but also for their conge-
nial and knowledgeable conversations. Harry captivated customers with his
storehouse of botanical and bibliographical knowledge. He and his wife regularly
attended the annual meetings of the Council on Botanical and Horticul-
tural Libraries, Inc., and national and international book fairs. Concerning
the latter, he attended 49 annual exhibits of the Frankfurt Book Fair in Germany.

Harry was a modest and caring person, yet he forthrightly and candidly ex-
pressed his opinions. I remember him also for many reminiscences of his book
trade career and the botanists with whom he associated. With a rich background
of experience and strong sense of the book market, Harry successfully intro-
duced the quality paperback, also known as the softbound, to the scholarly book
trade. Among the first in his field, he adopted photo-oftset printing as the pro-
cess for producing reprints. Even when this was not widely practiced, Harry
encouraged minority hiring and developed a staff fluent in numerous languages.
He held membership in the American Antiquarian Booksellers’ Association,
American Institute of Biological Sciences (AIBS), Council on Botanical and
Horticultural Libraries, Inc., International Association for Plant Taxonomy,
Mycological Society of America, New York Botanical Garden, North Ameri-
can Mycological Association, and Old Book Table (NY). Harry became a U.S.
citizen on 14 December 1936. As a member of the Lutheran Church, he was a
deacon during the 1960s and early 1970s. Among his honors was a Special Service
Award from the Mycological Society of America presented at the annual meet-
ing of the AIBS, San Antonio, TX, August 1991.

Sipa 18(1): 375. 1998

376 Stipa 18(1)

Harry’s successful life was shared with his wife Anne Johanne Marie Ficke)
whom he married on 2 October 1937. Harry is survived by his wife Anne;
two sons, Peter Thomas Lubrecht, Sr., a retired teacher of Drama and En-
glish and presently Artistic Director of Brundage Park Playhouse, of Randolph,
NJ, and Charles Frederick Lubrecht, co-manager of Lubrecht & Cramer, of
Montgomery, NY; three grandsons: Peter T. Lubrecht, Jr., Christopher C.
Lubrecht, and Bryan Leone. Harry's two sisters, Ilse and Ruth Lubrecht,
predeceased him. Funeral services were held 15 November 1997 in St. Peter’s
Lutheran Church, Port Jervis and cremation took place at the H. G. Smith
Crematory, Stroudsburg, PA. Interment of his ashes will take place at the
Forestburgh Cemetery. As a tribute to Harry, a memorial program in cel-
ebration of his life will be held at The New York Botanical Garden in the
fall 1998. Memorial contributions may be sent to The LuEsther T. Mertz
Library, New York Botanical Garden, 200th Street and Southern Boule-
vard, Bronx, NY 10458.

ACKNOWLEDGMENTS

I am grateful for information on Harry that Anne, Charles, and Peter
Lubrecht provided. Ronald L. Stuckey suggested this biographical essay and
supplied the photograph. The aforementioned individuals and Rogers McVaugh
read the manuscript and made constructive suggestions for changes in the
text. Additional information on Harry was obtained from a feature column
on bookmen (C[{hernofsky} 1986).

REFERENCES
Anonymous. 1952 Pe of-Print Department expands. Stechert-Hafner Book News 6(6):88.
1960. First Antiquarian Book Fair in America. Antiquarian Bookman

.1967.E Peli American botany catalog. Stechert-Hafner Book News 22(1): 6.
1972. 100 years {of} serving libraries. Stechert-Hafner Book News 26(5):41—43
Ctheenotsky), y. L. 1986. Lubrecht spans two eras of bookselling history. AB Hogkinen's S
78(5):357-358
Ewan, J eas 1967. Classica poetics Americana, pp. 38-40, In {H. Lubrecht}, Early American
otanical Works with a Miscellany of Other Botanical Rarities. Stechert-Hafner, Inc
New York.

*\9

- 1969. A Short History of Botany in the United States. Hafner Publishing
, New York ee ea

tubreche, H.}. ; mee American Botanical a with a Miscellany of Other Bo-

tanical eae Ae nc.,
. 1986. Ament of a rare- eee in Europe. AB Bookman’s Weekly
78(1 0): 808, 810

Stuessy, T.P., R.L. Studkey WL. aan araet and W.R. Burk. 1997. Botanical libraries
and herbaria in North America. 2. The Rudolph patel a library and its acquisi-
tion by The Ohio State University. Taxon 46:643—648

—_

Sipa 18(1): 376. 1998

First report of the genus Burmeistera (Campanulaceae) from Honduras

Thomas G. Lammers and P.J.M. Maas

363

Triraphis mollis (Poaceae: Arundineae) a species reported new to the United States
Stepban L. Hatch, William E. Fox Ill, John E. Dawson HI, and William B. Godwin

305

Priority of the name Agalinis harperi (Scrophulariaceae) over the names Agalinis delicatula
and Agalinis pinetorum

John F. Hays

369

In Memoriam—Heinz Dietrich Lubrecht (1908-1997), botanical bookseller and friend
William R. Burk
oye

Book notices and reviews 192, 236, 240, 246

Index of new names and new combinations in this issue

Agave gracilis Garcia-Mend. & E. Martinez, sp. nov. 227

Corsia purpurata var. wiakabui Takeuchi & Pipoly, var. nov. 164
Crataegus okanagensis J.B. Phipps & O’Kennon, sp. nov. 178
Crataegus okennonii J.B. Phipps, sp. nov.

Crataegus series Puspiccofrucd’): B. Phipps & O’Kennon, ser. nov. 184

ke fosteri Pipoly, sp. nov. 14
Cybianthus grandezii Pipoly, sp. nov. 139
ianthus granulosus Pipoly, sp. nov. 13
Cybianthus guyanensis subsp. pecudsicacorcis (Mig. in Mart.) Pipoly, comb. et stat. nov. 49
Cybianthus huampamiensis Pipoly, sp. nov. 130

y nso 142
cybiathus kayapii jac ) oh comb. nov. 84
Cybianthus nestorii Pipoly, sp. n

eybiantius Pe udolongifolius ra sp. nov. 101

n Pi 53

no
Mandevilla pringlei J.K. Williams, sp. nov. 231
Mandevilla holosericea (Sessé & Mog.) J.K. Williams, comb. nov. 237
Psychotria osiana Takeuchi & Pipoly, sp. nov. 162
Ruellia jimulcensis Villarrea nov
Sporobolus pinetorum Weakley & PM. Peterson, sp. nov. 258
Trachypogon mayaénsis Wiptt & S.D. Jones, sp. nov.
Xanthostemon fruticosus Peter G. Wilson & L. Co, sp. nov. 283

EG

Seasonal changes in concentration and distribution of heavy metals in creosotebush,
Larrea tridentata (Zygophyllaceae), tissues in the E] Paso, TX/Ciudad Juarez, Mexico area
William P. Mackay, Richard Mena, Nicholas E. Pingitore Jr., Keith Redetzke,

C. Edward Freeman, Harold Newman, John Gardea, and Hector Navarro

New plant records for Dominica, Lesser Antilles

Steven R. Hill and Arlington James

297

A new adder’s-tongue (Ophioglossum: Ophioglossaceae) for North America
James C. Zech, Patricia R. Manning, and Warren Herb Wagner Jr.

Notes on the flora of Texas with additions and other significant records
Larry E. Brown and Stuart J. Marcus
515

Cyperus fuscus (Cyperaceae), new to Missouri and Nevada, with comments

on its occurrence in North America

Paul M. McKenzie, Brad Jacobs, Charles T. Bryson, Gordon C. Tucker, and Richard Carter
325

The use of animal-dispersed seeds and fruits in forensic botany
Barney L. Lipscomb and George M. Diggs, Jr.
335

NOTES

Glaucium corniculatum (Papaveraceae) in Texas

Z0e Kirkpatrick and J.K. Williams

347

Hovenia dulcis (Rhamnaceae) naturalized in central Texas
Douglas H. Goldman

450

Habranthus tubispathus (Liliaceae) new to the flora of Alabama
Alvin R. Diamond, Jr., Charles P. Chapman, and Jim Brummett

Plantago coronopus (Plantaginaceae) new to Texas
Robert J. O’Kennon, George M. Diggs, Jr. and Ronald K. Hoggard
356

Pteris vittata (Pteridaceae), a new fern for Texas

Jack W. Stanford and George M. Diggs, Jr.

SP,

Lycianthes asarifolia (Solanaceae) new and weedy in Texas
Monique Dubrule Reed and Mary Ketchersid

301

a N (continued on inside back cover)

BRIT ISSN 0036-1488

CONTENTS

Arnoglossum album (Asteraceae): New species from northern Florida

Loran C. Anderson

377

Seven new nomencl ature il COIL mbinat tions and a new name in Packera ( (Asteraceae: Senecioneac)
Debra K. Trock and Theodore M. Barkley

385

Ardisia niambiensis (Myrsinaceac), a new species of Ardisia subgenus Ardisia from the Chocé
Floristic Province of Colombia

Jobn fj. Pipoly Hand Alvaro Cogollo P

389

Clusia niambiensis (Clusiaccac), a new species from the Choco Floristic Province of Colombia and
Ecuador

John J. Pipoly If, Alvaro Cogollo P., and Marta Sofia Gonzélez

305

notes in Clusia (Clusiaceae) from Andean Colombia and Venezuela

Jobn J. Pipoly Ill and Alvaro Cogollo P
401

New species and nomenclatura

Dos nuevas adiciones a la orquideoflora Mexicana
Adolfo Espejo Serna, Ana Rosa Lopez-Ferrari, Javier Garcia Cruz, Rolando Jiménez Machorro y Luis
Sainchez Saldana

tH]
seedling development in species of Chamaesyce (Euphorbiaceae) with erect growth habits

W. John Hayden and Olga Troyanskaya

419

A revision of the genus Ardisia subgenus Graphardisia (Myrsinaceae)
John J. Pipoly Ill and Jon M. Ricketson

433

Una nueva epee es Polianthes (Agavaceae) del estado de Ouxaca, México
Eloy Sola ind Abisai Garcia-Mendoza

473

Pollen morphology of the genus Echinopepon (Cucurbitaceae)
Concepcion Rodriguez J. and Rodolfo Palacios-Chdvez
479

Variation in the Berlandiera pumila (Asteraceae) complex
G.L. Nesom and B.L. Turner

New names and combinations in neotropical Myrsinaccue
Jobn J-Pipoly I and Jon M. Ricketson
503

A new species of filmy fem (Aymenophyllaceae: Pteridophyta) froni South India
. Abdul Hameed and PY. Madbusoodanan
519
A second species of Oritrophium (Asteraceae: Astereac) from Mexico
Guy L. Nesom ree ea ie
ans CONTRIBUTIONS
923
: | TO BOTANY
Uniqueness of the endangered Florida Semaphore Cactus (Opuntia corallicola)
Daniel F. Austin, David M. Binninger, and Damald J, Pirikava
527

Arkansas Carex (Cyperaceae): A briefly annotited list
Philip E. Hya VOLUME 18
pe NUMBER 2

(continued on back cover) DECEMBER, 1998

CONTRIBUTIONS TO BOTANY

FOUNDED BY
LLOYD H. SHINNERS
1962

we

Wm. F. Mahler
Publisher 1971-1992

Ww

5)
G

Barney L. Lipscomb

Editor

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Fort Worth, Texas 70102-4000, USA

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Electronic mail: sida@brit.org
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Dpto. de Botanica, Facultad de Biologia
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Contributing Spanish Editor

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E-24071 Leon, Spain

Guidelines for contributors are available upon request
and on the inside back cover of the last issue of each volume.
Subscription per year: $25. Individual, $45. USA Institutions, $5

0. Outside USA;
numbers issued twice a year

© SIDA, CONTRIBUTIONS TO BOTANY, Volume 18, Number 2, pages 377-630.
Copyright 1998
Botanical Research Institute of Texas, Ine,
Printed in the United States of America
|

SSN 0030-1488

ARNOGLOSSUM ALBUM (ASTERACEAE): NEW
SPECIES FROM NORTHERN FLORIDA

LORAN C. ANDERSON

Department of Biological Science
Florida State University
Tallahassee, FL 32306-4370, U.S.A.

ABSTRACT
The new species Arnoglossum album is described. Its phyllaries have prominently winged
keels—a feature shared with A. diversifolium, A. floridanum, A. pl antagineum, and A. sulcatum.
Comparisons among members of this closely knit assemblage include uae
nology, ecological setting, and range, and a key | is provided to distinguish the spec

RESUMEN

Se describe una especie nueva Arnoglossum album. Sus filarios tienen quillas aladas
prominentes—una caracterfstica que comparte con A. ge A. ee eonate A
plantagineum, y A. sulcatum. |
reticulado incluye morfologia, Fenologi caracteristicas ecoldégicas y are mal. Se oftece una

clave para diferenciar las espec

Species of the tussilaginoid Armoglossum, commonly called “Indian Plantains,”
were formerly placed in the heterogeneous “Cacalia” (Kral & Godfrey 1958;
Cronquist 1978; Pippen 1978), but, because Cacalia is now typified by a
species of the senecionoid European genus “Adenostyles” and because ge-
neric circumscriptions are now more narrowly drawn, Arnoglossum is con-
sidered distinct by many recent authors (Robinson 1974, 1980; Jeffrey 1979,
1992; Godfrey & Wooten 1981; Wetter 1983; Funk 1985; Brummitt 1992).
Arnoglossum has three species with extensive ranges in the eastern United
States [A. atriplicifolium (L.) H. Robins. and A. reniforme (Hook.) H. Robins. ]
or southeastern United States [A. ovatum (Walt.) H. Robins.] and one from
the central part of the country [A. plantagineum Raf.]. The remaining three
species have more limited ranges, one as a Florida endemic [A. floridanum
(A. Gray) H. Robins.] and the others with ranges barely extending from
Florida into neighboring states [A. diversifolium (Torr. & A. Gray) H. Robins.
and A. sulcatum (Fern.) H. Robins.]. The latter four species (.e., A. plantagineum
and “the Florida group”) are ata in having phyllaries with keels that are
strongly winged. The new species d d below is now added to this assemblage
with winged phyllaries.

Sipa 18(2): 377-384. 1998

378 Sipa 18(2)
MATERIALS AND METHODS

Fresh and dried materials were processed as described by Anderson (1994).
Heads from at least five different collections per species were dissected and
measured for involucral and floral quantitative data. Living plants of the
five species under study were propagated from seed or as transplants in a
greenhouse or common garden in Tallahassee, Florida. Vouchers of seed-
lings, plants at various stages of development, and all cited specimens are
deposited at FSU unless noted otherwise.

SPECIES DESCRIPTION
oe album L.C. Anderson, sp. nov. (Figs. 1, 6-9). Type: U.S.A.
LORIDA. Bay Co.: locally common in acidic, poorly adned sandy soil of open,
wet savanna with Sarracenia flava, Rhynchospora, Rhexia, and cas 7 Spp. E of
Rte 77 on N side of Southport, T2S, R14W, NE1/4 of SE1/4 Sec 21, 6 Jun 1995,
L.C. Anderson 15555 (HOLOTYPE: NY!; ISOTYPES: BRIT! a “MO! Us).

Herbae perennes plerumque 8-10 dm alta. Folia basalia 20-63 cm longa, S—16 cm lata,

longe one laminis late ovatis vel anguste oblongo-lanceolatis, vernis lateralibus orincipalibus

ra basin laminae ad venam mediam arcte parallelis dein divergentibus. Folia
caulina pauca, ee 26 cm longa, petiolata, serrata, foliis superioribus reductis sessilibusque.
Capitula in cymis corymbosis disposita, involucris 10-13.4 mm longis, phyllariis albis carinis
prominenter alatis. Corollae albae, 9-10.5 mm longae.

Essentially glabrous cespitose herbs. Stems (5—)8—10(—10.5) dm tall, light
green, strongly ridged. Basal (radical) leaves (20—)30—55(—63) cm long overall,
5—10(—16) cm wide, long petiolate, (10—)15—18(—34) cm, blades broadly
ovate to narrowly oblong-lanceolate, margins entire, shallowly sinuate, or
rarely serrulate-denticulate, bases attenuate, apices rounded, mucronulate,
basally disposed lateral veins appressed to the midrib 2—4 cm then abruptly
extending toward the leaf margins; cauline leaves few, blades (7—)12—18(—
26) cm long overall, (2.5—)4—5(—6) cm wide, petioles (1-)4—5(-12) cm long,
ovate, bases cuneate, apices acuminate, serrately toothed, upper leaves re-
duced in size and sessile. Inflorescence compound, freely branched, + flat-
topped, with heads crowded in corymbose cymes. Heads 5-flowered; in-
volucres 10—12.5(-13.4) mm long, cylindric, phyllaries 5, chalky white,
margins hyaline, keels prominently winged (the wings rising 1—-1.5(—2.0)
mm from the phyllaries with margins sinuate to erose and reduced apically).
Receptacles with short central cusp. Corollas white, rarely tinged with pink,
9—-10.5 mm long, veins 10, tube 5—6.4 mm, throat 0.2—1.0 mm, lobes 2.9—
3.8 mm, lanceolate; anthers 2.5—2.6 mm long, collars cylindrical; styles 11.8—
12.5 mm long, style branches 1.5-1.7 mm long, stylopodium immersed
in nectary; achenes glabrous, 4.2-5 mm long, cylindric to narrowly clav-
ate, carpopodium cells erect, pappus white, 6-7 mm long. » = 25.

Distribution.—Endemic to Bay and Gulf counties of Florida in poorly

ANDERSON, A new species of Arnoglossum

a7)

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Fics. 1-24. Representative leaves of selected Arnoglossum species, all at 1/6 full size. Figs.
1-5. Radical leaves showing ae ate A. album with appressed lateral veins,
Anderson 15600, Bay Co., FL. Fig. 2 pe Anderson 13414, Levy Co., FL.
3. A. floridanum, Anderson 12165, ‘Tavlos Co., FL.
Riley Co., KS. Fig. 5.

Fig.
Fig. 4. A. plantginenn, Anes 12] Ce

A. sulcatum, Anderson 16035, Santa Rosa Co. igs. 6—24. Out-
lines of cauline leaves (major venation similar to respective pidical ee ae 6-9. A.
album. Fig. 6. Anderson 12137, Gulf Co., FL. Figs. 7-8. Anderson 15600, Bay Co., FL.
Fig. 9. Anderson 13432, Gulf Co., FL. Figs. 10- 13 : doagilinn Fig. 10. Anderson 13475,
Levy Co., FL. Figs. 11, 13. Anderson 13478, Putnam Co., FL. Fig. 12. McDaniel 9059,
Houston Co., AL. Figs. 14-15. A. floridanum. Fig. 14. Anderson 12160, Taylor Co., FL.
Figs. 15-16. Anderson 12077, Clay Co., FL. Figs. 17-20. A. es ie Fig. 17. Mehrhoff
12299 (CONN), Bruce Co., Ontario Prov., Canada. Fig. 1 n 11991, Loundes
Co., AL. Fig. 19. Bryson 9861, Oktibbeha Co., MS. Fig. 20. ee. 11968, Tangipohoa
Par., LA. Figs. 21-24. A. sulcatum. Fig. 21. McDaniel 7978, Geneva Co., AL. Fig. 22. Anderson

12342, Leon Co., FL. Fig. 23. Anderson 16058, Walton Co., FL. Fig. 24. Anderson 16057,
Walton Co., FL.

380 Sipa 18(Q)

drained, acidic (pH 5-6), loamy sands (with 0.35—1.7% organic matter)
of wet savannas, open pinewoods, and most frequently the ecotone between
the two, often in close association with several of the following: Aletris lutea,
Aristida beyrichiana, Asclepias longifolia, A. michauxti, A. viridula, Aster
eryneiifolius, Cleistes bifaria, Drosera capillaris, Eriocaulon compressum, Hy-
pericum gymnanthum, Lachnanthes caroliniana, Lachnocaulon anceps, Lo-
belia floridana, Ludwigia linifolia, L. microcarpa, L. virgata, Marshallia tenuifolia,
Oxypolis greenmanit, Panicum rigidulum, Physostegia godfreyi, Platanthera
nivea, Polygala balduinii, P. cruciata, P. ramosa, Rhexia lutea, Rhynchospora
ciliaris, R. curtisii, R. fascicularis, R. filifolia, R. globularis, R. oligantha, R.
pusilla, R. variflora, R. wrightiana, Rudbeckia graminifolia, Ruellia noctiflora,
Sabatia bartramii, S. campanulata, Sarracenia flava, Scutellaria floridana,
lofieldia racemosa, Verbesina chapmanii, Xyris ambigua, X. baldwiniana, and
X. elliotii. The woody associates (less frequent) include: Cyrilla racemiflora,
Hypericum cistifolium, H. exile, H. reductum, Hex coriacea, 1. vomitoria, Magnolia
virginiana, Myrica cerifera, Nyssa ursina, and Pinus elliottit. Flowers June to
mid-July.

Representative specimens ex: amined. U.S.A. FLORIDA. Bay Co.: type locality, Z.C. Anderson
15529 (FSU), 20 Jun 1995, L.C. Anderson 15600 (FSU, MO, NY), 20 Jun 1995, L.C.
Anderson 15601 (FSU), L.C. ‘Aan 15708 (FSU); S side County Rd 2300, 1.9 mi W of

5 air mt NW of Southport, 13 Jun 1995, £.C. Anderson 15577 (FSU); 0.5 mi E
of Burnt Mills Creek bridge on S side of Rte 388, 1.9 mi W of Rte 77, 1.5 air mi WNW
of Southport, 13 Jun 1995, L.C. Anderson 15581 (FSU). Gulf Co.: 5.3 mi S of Rte 22 on
E side of Daniels Rd near Wetappo Creek, ca. 5.5 air mi WSW of Wewa- hitchka, L.C.
Anderson 11642 (FSU), L.C. Anderson 11673 (2), L.C. Anderson 11934 (FSU), L.C. ii ne
12137 (BRIT, FSU, GH, MO, NY, US), £.C. Anderson 12138 (FSU), L.C. Anderson 1342
(FSU), 5. at mi S of Rte 22 on W side Daniels Rd, 1.C. Anderson 11950 (FSU), L.C. eye
12171 (FS

DISCUSSION

The Indian plantains with winged phyllaries form a closely related alli-
ance of species. [hey differ from each other in several floral features as summarized
in Table 1. The longest involucres occur in A. floridanum, and the shortest
in A. sulcatum. Phyllaries in A. album have prominent wings proximally
that become highly reduced apically, whereas in all other species the wings
are shorter in height but + equally developed for the length of the bract
(often broadest apically). Arnoglossum album also has the whitest phyllar-
ies; in the other species they are more greenish-white.

Corollas of A. album are white, rarely tinged with pink; in the other species
they are white, light lavender, or ochroleucous (more greenish-white or yellowish-
white in A. floridanum and sometimes more pinkish in A. diversifolium).
Corollas are generally longest in A. floridanum and shortest in A. suleatum,
but not as short as reported by Pippen (1978) and Cronquist (1980). In all
species the corolla tubes are relatively long and the throats below the lobes

ANDERSON, A new species of Arnoglossum 381

TABLE 1. Size floral features (in mm) of “winged” Arnoglossum species.

‘Taxon Involucre length Corolla length Tube width Lobe length
. album .0-13.4 9.0-10.5 0.9-1.0 2.9-3.8

A. diversifolium 10.2 2-11.0 9.1-10.2 0.8-0.9 3.1-3.5

A. floridanum .0-14.8 9.8—12.2 0.8—0.9 2.84.1

A. plantagineum eee 9.2-11.4 0.7-0.8 3.1-3.6

A. sulcatum 8.5-9.9 7 .8-9.6 0.6—0.7 2.7-3.4

very short. The tube widths are greatest in A. album and most slender in A.
sulcatum. Carpopodial cells of the achenes in A. album are generally erect,
whereas they are procumbent in A. plantagineum. Cell shape varies some-
what within a given population because of differing stages of achene matu-
rity. Robinson and Brettell (1973) suggested carpopodial structure was an
excellent character for distinguishing genera in the Senecioneae; they re-
corded procumbent carpopodial cells for Mesadenia (i.e., Arnoglossum). Wetter
(1983) found considerable variation in this and other micromorphological
characters and questioned their value as generic markers.

Vegetatively, A. album looks most like A. plantagineum, and that species
is probably its closest relative. The two differ in seedling morphology. Cotyledons
of A. album are 1-1.5 cm long and 9-12 mm wide (+ orbicular), whereas
they are 3-4.5 cm long and 8-9 mm wide (spatulate-oblanceolate) in A.
plantagineum.

The five species differ more in leaf morphology than they do in floral
features (Figs. 1-24). Basal leaves of A. album have distinctive venation.
The principal lateral veins are appressed and closely parallel the midvein
into the blade for a few centimeters and then spread abruptly toward the
leaf margins (Fig. 1). This feature of appressed lateral veins is also found in
the related Yermo xanthocephalus of Wyoming. In the other Arnoglossum species,
the lateral veins spread immediately at the base of the blade (Figs. 2-5).

Relative length of the petiole on radical leaves is often used in keys to
the species (Kral & Godfrey 1958; Pippen 1978; Cronquist 1980). This
feature is correlated to habitat y of the species. Generally, A. diversifolium

and A. sulcatum grow in deep shade, and their radical leaves have long peti-

oles (Figs. 2, 5). Arnoglossum floridanum usually grows in full sun, and its
radical leaves have short petioles (Fig. 3). Petiole lengths are more variable
in radical leaves of A. album and A. plantagineum. These two species often
grow in savannas or prairies. Their petiole lengths are related to the density
of grass and other vegetation near the plants; plants from the same seed
source that were grown in different shade or vegetation densities in my garden
produced short-petioled leaves in sunnier or less crowded situations and
longer petioles in shaded or crowded conditions.

382 Sipa 18(2)

Cauline leaves of A. diversifolium (Figs. 10-13) are petiolate and basally
truncated or cordate; they are petiolate and rounded to broadly cuneate in
A. album (Figs. 6-9), A. floridanum (Figs. 14-16), and A. plantagineum
(Figs. 17-20), whereas they are + sessile and more narrowly cuneate in A.
sulcatum (Figs. 21-24). Cauline leaves of A. diversifolium are deltoid-hastate
in outline and dentately toothed. They tend to be ovate, serrately lobed,
and acute in A. album; ovate-oblong, crenulate, and obtuse or rounded in
A. floridanum; ovate to lanceolate or narrowly spatulate, entire or serru-
late, and acute in A. plantagineum;, and ovate to rhombic or narrowly lan-
ceolate, sparsely serrate-dentate, and acute to obtuse in A. sulcatum.

Arnoglossum plantagineum has the greatest geographical range among the
five species (east Texas, Louisiana, and the black soil belt of Alabama north
through Kansas and Ohio to southern Minnesota, Michigan, and Ontario,
Canada); it occurs mainly in calcareous, tall-grass prairie (Pippen & Chapman
1986), but minor habitats include glades, fens, and pine-oak woodlands.
Arnoglossum floridanum occurs in well-drained sands in open pine-scrub
oak, dry flatwoods, and old fields from Duval and Madison through pen-
insular Florida south to Highlands and Manatee counties.

The other three species occur mainly in the Florida panhandle. Arnoglossum
diverstfolium occurs in river swamps and wet hammocks and ts the most
wide-ranging of these three; it has a few populations in southern Alabama
and southwestern Georgia, ranges from Walton County to Leon County
in the panhandle, and has disjunct populations in Putnam, Volusia, and
Levy counties, Florida. Arnoglossim sulcatum occupies shaded acid bogs or
swamps; it occurs sporadically in extreme southern Alabama and south-
western Georgia and from Escambia to Leon Counties of the Florida pan-
handle. Arnoglossum album is restricted to Bay and Gulf Counties of the
Florida panhandle; the only other species of the group that occurs in those
counties ts A. sulcatum, but it has different habitat preferences and
phenology.

Flowering time in Arnoglossum plantagineum is clinal from south to north;
the plants blooming in late April in southern Louisiana to early August in
Minnesota and Michigan (Pippen & Chapman 1986). A reverse cline ex-
ists for A. ee it blooms in the Florida panhandle from May to

early July (same period for A. floridanum), whereas, to the south, in Levy
County (where it is sympatric with Hasteola robertiorum) A. diversifolium
blooms from mid-August through September. Arnoglossum album blooms
from June to mid-July, and A. sudcatum is the latest to bloom (September
to October).
he new species, A. album, is distinguished from all of its immediate
relatives by the following features: it is geographically isolated from all other
species except A. sulcatum, and it is totally separated from A. sulcatum by

ANDERSON, A new species of Arnoglossum 383

habitat and phenology; basal leaf venation is distinctive; cauline leaves are
most similar to those of A. sulcatum, but they are generally narrower and
sessile in A. sudcatum; its involucres and corollas appear whiter than those
of the other species; the wings of the phyllaries are higher proximally and
attenuated and distally, whereas wings are lower in height overall and evenly
raised along the keel or somewhat higher distally in the others; the phyl-
lary wings have erose margins distally in A. album, and wing margins are
entire or sinuate in the other species; and the corolla tubes are wider than
those in any other species in the group.

KEY TO THE FIVE SPECIES OF ARNOGLOSSUM WITH WINGED PHYLLARIES

Most keys to species of Arnoglossum include significant habitat and phe-
nological data. The following key uses only morphological features.

1. Blades of radical leaves truncate, ovate to cordate-ovate, sparsely den-
ticulate; lower cauline leaves deltoid-hastate, dentately toothed........ A. diversifolium
1. Blades of radical leaves ovate to ovate-oblong, entire, sinuate or crenate;
lower cauline leaves ovate, entire, crenulate or serrately toothed.
2. Phyllaries with prominently winged keels, wings highest at base, erose;
radical leaves with lateral veins appressed to midveins for 2—4 c
then spreading. A. album
; Phyllaries with keels + evenly winged throughout or highest apically,
entire or sinuate; radical leaves with lateral veins diverging from base
of blade.
3. Involucres and corollas mostly over 11 mm and 10 mm long, re-

Pee nine leaves crenulate A. floridanum
] 1 ] .

iw)

ula
4, Involucres mostly over 10 mm long, corollas over oc mm; seanlin

leaves at midstem rounded to petiolate base. A. plantagineum
4, Involucres less than 10 mm long, corollas 8—9(—9.6) mm; cauline
leaves at midstem broadly cuneate and sessile A. sulcatum

ACKNOWLEDGMENTS

This study was supported in part by a grant from the Council on Re-
search and Creativity of Florida State University. Richard Pippen graciously
shared his knowledge of the group. Mark Garland assisted with the Latin
diagnosis, and Ken Womble helped with the illustrations. Ted M. Barkley,
R.R. Kowal and A.B. Thistle are thanked for providing critical reviews of

the manuscript.

REFERENCES
ANDERSON, L.C. 1994. A revision of Hasteola (Asteraceae) in the New World. Syst. Bot.
19:211-
Brummitt, R.K. 1992. Vascular plant families and genera. Kew: Royal as ee
Cronquist, A. 1978. Compositae, tribe Senecioneae. N. Amer. Flora, ser. I], 10:14-16.
1980. Vascular plants of the southeastern United States. Vol. 1. aes Univ.

384 Sipa 18(2)
of North Carolina Press, Chapel Hill.
Funk, V.A. 1985. Cladistics and generic concepts in the Compositae. Taxon 34:72-80.
Goprrey, R.K., and J.W. Wooren. 1981. Aquatic and wetland plants of the southeastern
United States. Dicotyledons. Univ. of Georgia Press, Athens.
Jerrrey, C. 1979. Generic and sectional limits in Senecio (Compositae): IH. Evaluation of
some recent studies. Kew Bull. 34:49-58
. 1992. The tribe Senecioneae (C jae in the Mascarene “red with an an-
notated world check-list of the genera of the tribe. Kew Bull. 74:49
Krat, R., and R.K. GODEREY. ete rae of the Florida species of aL ee sompositae).
Quart. J. Florida Acad. Sci. 21:193-206.
Pippen, R.W. 1978. Cacalia. N. ree Flora, series [1], 10:151—159.
and K.A. CHAPMAN. 1986. Comparison of morphological characters between re-
gional habitats of Cacalia plantaginea (Asteraceae). In: G.K. Clanby and PN. Pemble,
eds. Proceedings of the 9th North American Prairie C ie tieies ‘Tri-college University
Center for Environmental Studies, Fargo,
Rosinson, H. 1974. ee lathe Seneeianese neingeds:
Phytologia 28:294—2¢
. 1980. ee Tse a (Hook.) H.Robins., comb. nov. Phytologia 46:441,
and R.D. Brettell. 1973. Studies in the Senecioneae (Asteraceae). IV. The genera
Mesadenia, Syneilesis, Hea Koyamacalia and Sinacalia. Phytologia 27:265-276.
Werrer, M.A. 1983. Micromorphological characters and generic delimitation of some
New World Senecioneae (Asteraceae). Brittonia 35;1—22

Nae

VI. The genus Arnoglossum.
ec c

SEVEN NEW NOMENCLATURAL
COMBINATIONS AND A NEW NAME IN
PACKERA (ASTERACEAE: SENECIONEAE)

DEBRA K. TROCK and THEODORE M. BARKLEY!

Herbarium, Division of Biology
Kansas State University
Manhattan, KS 66506, U.S.A.
dkwel@ksu.edu

ABSTRACT

The following new combinations and new name are proposed: Packera bolanderi var.
harfordii (Greenm.) D.K. Trock & T.M. Barkley, comb. nov.; Packera dimorphophylla vat.
intermedia (T.M. Barkley) D.K. Trock & T.M. Barkley, comb. nov.; Packera dimorphophylla
var. paysont (1.M. Barkley) D.K. Trock & T.M. Barkley, comb. nov.; Packera pseudaurea
var. flavula (Greene) D.K. Trock &T.M. Barkley, comb. nov.; Packera pseudaurea var. semicordata
(Ma ck. & Bu sh ) D.K. ‘Tro ck & TM. Barkley, comb. NOV.; £ Packera neomexicana vat. toumeyl
(Greene) D.K. Trock & T.M. Barkley, comb. nov.; Packera neomexicana var. metcalfei (Greene)
D.K. Trock & T.M. Barkley, comb. nov.; and Packera buekii D.K. Trock & T.M. Barkley,

nom. Nov.

RESUMEN

Se proponen las siguientes nuevas combinaciones y nuevo nombre: Packera bolanderi
var. harfordii (Greenm.) D.K. Trock & T.M. Barkley, comb. nov.; Packera dimorphophylla
var. intermedia (T.M. Barkley) D.K. Trock & TM. Barkley, comb. nov.; Packera dimorphophylla
var. paysont (T.M. Barkley) D.K. Trock & T.M. Barkley, comb. nov.; Packera pseudaurea
var. flavula (Greene) D.K. Trock & TM. Barkley, comb. nov.; Packera yon ie semicordata
(Mack. & Bush) D.K. Trock & T.M. Barkley, comb. nov.; ere neomexicana var. toumeyi
(Greene) D.K. Trock & T.M. Barkley, comb. nov.; Packera neomexicana var. metcalfei (Greene)
D.K. Trock & T.M. Barkley, comb. nov.; y Packera buekii D.K. Trock & T.M. Barkley,

nom. nov.

The genus Packera (Asteraceae: Senecioneae) has been known widely as
the Aureoid group of Senecio, and the recognition of Packera necessitates
many nomenclatural transfers. Preparation of the treatment of Packera for
the Flora of North America project has drawn our attention to the trans-
fers proposed here. Type specimens for the basionyms are provided by Barkley
(1978). The combinations P pseudaurea var. semicordata and P dimorphophylla
var. intermedia are proposed simply for consistency in the use of “varietas”
'Present address: Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX
76102-4060, U.S.A, e-mail: barkley@brit.org

SIDA 18(2): 385-387. 1998

386 Stipa 18(2)

as the infraspecific rank of choice in Packera The two epithets are other-
wise available as subspecies.

Packera bolanderi var. harfordii (Greenm.) D.K. Trock & T.M. Barkley,
comb.nov. BASIONYM: Senecio harfordii Greenm. in Piper, Contr. U.S. Natl. Herb.
11:567. 1906. Senecio bolanderi var. harfordii (Greenm.) V.M. Barkley

Packera dimorphophylla var. intermedia (1.M. Barkley) D.K. Trock & T.M.

Barkley, comb. nov. BASIONYM: Senecio dimorphophyllus var. intermedius I.M.
Barkley, Trans. Kansas Acad. Sci. 65:362. 1963. Packera dimorphophylla subsp. intermedia
(ILM. Barkley) Weber & Love.
Packera dimorphophylla var. paysoni (1.M. Barkley) D.K. Trock & TM.
ar ley, comb. nov. — ye necio dimorphophyllus var. paysoni 1M. Barkley,
‘Trans. Kansas Acad. Sci. 65:362.

Packera pseudaurea var. flavula ——o D.K. Trock & T.M. Barkley, comb.
nov. BASIONYM: Senecio flavulus Greene, Pittonia 4:108. 1900. Senecio pseudaureus
var. flavula (Greene) Greenm. Packera pseudaurea su bsp. flavula (Greene) Weber &
Léve.

Packera pseudaurea var. semicordata (Mack & Bush) D.K. Trock & T.M.
Barkley, comb. nov. BASIONYM: Senecio semicordatus Mack. & Bush, Rep.
Missouri Bot. Gard. 16:107. 1905. Senecio aureus var. semicordatus (Mack. & Bush)
Greenm. Senecio pseudaureus var. semicordatus (Mack. & Bush) T.M. Barkley.

Packera neomexicana var. toumeyi (Greene) D.K. ‘Trock & T.M. Barkley,
comb. nov. BASIONYM: Senecio daca eniti Pittonia 3:349. 1898. Senecio neo-
mexicanus var. toumeyi (Greene) T.M. Barkley

Packera neomexicana var. metcalfei (Cd D.K. Trock & T.M. Barkley,
comb. nov. BASIONYM: Senecio metcalfei Greene ex Wooton & Standl., Contr. U.S.
Natl. Herb. 16:193. 1913. Senecio neomexicanus var. metcalfei (Greene) T.M. Barkley.

The situation with the epithet “cymbalarioides” is complicated. A single-
headed plant of wet, open sites in western U.S. and adjacent Canada was
long known as Senecio subnudus DC., a name that was introduced by A.P.
de Candolle in 1837. Unfortunately, de Candolle used the epithet “subnudus”
twice; the other time for an African plant. Heinrich W. Buek (1840) rec-
ognized the error in his index to the Prodromus and he chose the African
plant to be the correct Senecio subnudus. He then called the American plant
Senecio cymbalartoides A. Buek, and unfortunately, this name remained forgotten
until the 1960s (Barkley 1962). In the meantime, a widespread plant of
western North America was named Senecio cymbalarioides by Thomas Nuttall.
It was necessary to change the name of the North American S. subnudus
DC. tes. cymbalarioides H. Buek, and S. cymbalarioides Nutt. then be-
came S. streptanthifolius Greene, which is the next oldest available name
for that entity (Barkley 1962). Recognition of the genus Packera required

TROCK AND BARKLEY, New combinations and a new name in Packera 387

the transfer of many epithets into that genus and, unfortunately, Senecio
cymbatarioides Nutt. was ferred into Packera where it became 2 cymbalarioides
(Nutt.) W.A. Weber & A. Love, (Weber & Léve 1981). Shortly thereafter,
Weber and Love recognized that it was S. cymbalarioides H. Buek that should
have been transferred to Packera, so they made the combination: P cymbalaroides
(H. Buek) W.A. Weber & A. Love (Weber 1984). The transfer of the Nuttall
epithet into Packera preoccupies that name in Packera so Packera cymbalioides
(H. Buek) W.A. Weber & A. Léve is illegitimate and must be rejected ac-
cording the International Code of Botanical Nomenclature (Greuter et al.
1994). Thus, the American plant first known as Senecio subnudus DC. and
then as Senecio cymblalrioides Buck must have a new name. The new epi-
thet that we chose for this distinctive plant commemorates Heinrich Wilhelm
Buek, the 19th Century German botanist whose carefully prepared index
to de Candolle’s Prodromus gave rise to the need for this new name.
Packera buekii D.K. Trock & T.M. Barkley, nom. nov. penn Senecio subnudus
DC., Prodr. 6:37. 1837 (non S. hers DC.,, Prodr. 6:405. 18
Buek, Gen. Sp. Synon. Cand. 2:6. 1840 (non S. ee thes Tans. Amer.
Phil. Soc. II. 7:412.1841.).

ACKNOWLEDGMENTS

We thank John Bain, John Pipoly, Alan Whittemore, John Strother, K.
Gandhi, and D.H. Nicolson for their advice and editorial wisdom in the
preparation of this work. This is contribution no: 99-/44-/ from the Kan-
sas Agricultural Experiment Station, Manhattan.

REFERENCES
ve EY, I.M. 1962. revision of Senecio aureus Linn. and allied species. Trans. Kansas
cad. Sci. 65:318—4
1978. Senecio. i. ‘Nee American flora II. 10:50—137.

Burk, H.W. 1840. 0. Genera, species, et synonyma Candolleana. 2.

CANDOLLE, A.P. de, 1837. Prodromus systematis. 05.

GreuTter, W., ER. Barriz, H.M. Burbet, W.G. Gone: i. DEMOULIN, D.L. HawkswortH,
P.M. JOERGENSEN, D.H. Nicotson, P.C. Sitva, P. TEHANE, & J.M. MCNEILL, eds. Inter-
national code of botanical nomenclature (Tokyo Code). Regnum. Veg. 131:1-389

Weper, W.A. 1984. New names and combinations, principally in the Rocky Mountain
flora-IV. Phytologia 55:1

Weer, W.A. and A. Love. 1981. New combinations in the genus Packera (Asteraceae).
Division 49:44—-50).

Sipa 18(2)
BOOK REVIEWS

A. Espgjo and A.R. LOpez-Ferrari. 1997. Las Monocotiledéneas Mexicanas.

er Sinopsis Floristica 1. Lista de Referencia PARTE VIII. Orchidaceae

1. (ISBN 968-6144-10-2; 968-6144-20-X pbk). Consejo Nacional de

la Flora de México, Apartado Postal 70-261, Delegacion Coyoacan,

04510 México, D.E and Universidad Auténoma M Iztapalapa,

Av Michoacan y La Purisima, Col. Vicentina, Delegacién Iztapalpa,
09340 México, D.F. No price given. 90 pp. Color cover.

A. Esprjo and A.R. Lopez-Ferrari. 1997. 1998. Las Monocotiledéneas
Mexicanas. Una Sinopsis Floristica 1. Lista de Referencia PARTE
VIII. Orchidaceae 2. (ISBN 968-6144-10-2; 968-6144-21-8 pbk Consejo
Nacional de la Flora de México, Apartado Postal 70-261, Delegacién
Coyoacan, 04510 México, D.E and Universidad Auténoma Metropolitana-
Iztapalapa, Av Michoacan y La Purisima, Col. Vicentina, Delegacion
Iztapalpa, 09340 México, D.F. No price given. 115 pp. Color cover.

These two volumes list the species of Orchidaceae known from Mexico, in alphabetical order by ge-
nus, then by spe cals from rile ta tO fee Orchid aceae 1) and Hom Perea to Xylobium
i through oO inaceae),
lete li ication, followed by iba nee along with all taxonomic

(Orchidaceae 2)

For each s spec “le
and oom ets al synonyms wand respective ype citations. It is clear that the authors have seen the
vast majority of the type specimens involved. There are no lectotypes chosen in the work, but lecto-
types chosen by other authors are cited cle: ae : pe specimens seen have the traditional“
otherwise, Seca of the holotype is cited as it appears in the protologue. Each species’ distribution
within Mexico is also given,

The list is a very handy reference for anyone interested in the Orchid flora of Mexico, or for that

matter, Mesoamerica. I is printed on high quality paper and is clear and easy to read. The author
abbreviations edalaioa to eno of a Names, Laas by the Royal Botanic Gardens, Kew,
and the literature cit id to the B-P-H and TL-2. I have seen no notable typographical
errors,

Annotated checklists such as these are extremely useful, especially given the complete index to ac-

id synomyms in the back. This series is absolutely necessary for any curator dealing
with Mexican monocotyledonous plants. We anxiously await the arrival of the other volumes, and
unhesitatingly recommend them for everyone!—/ohn J. Pipoly II

Sipa 18(2): 388. 1998

ARDISTIA NIAMBIENSIS (MYRSINACEAE),

A NEW SPECIES OF ARD/SIA SUBGENUS

ARDISIA FROM THE CHOCO FLORISTIC
PROVINCE OF COLOMBIA

JOHN J. PIPOLY III

Botanical Research Institute of Texas
509 Pecan Street, Fort Worth, TX 76102-4060 U.S.A.
Jpipoly@brit.org; clusia@latinmail.com

ALVARO COGOLLO P.

Fundacion Jardin Botdnico, Joaquin Antonio Uribe
Apartado Aéreo 51407, Medellin, Antioquia, COLOMBIA
acogollo @lati L.com; jardinbo @epm.net.co

ABSTRACT

Documentation of the flora of the Natural Reserve Rio Nambf has resulted in the dis-
covery of a heretofore undescribed species, Ardisia niambiensis. The species is described,
illustrated and its salient morphological features are elucidated. Ardisia niambiensis is best
placed in subgenus Ardisia because of its terminal, glandular-papillate, paniculate inflo-
rescences with tripinnate branches, and the evenly divided, 5-lobed calyx, whose symmet-
ric lobes have glandular-ciliolate margins. It is most closely related to Ardisia premontana,
a taxon endemic to the eastern slopes of the Eastern Andean Cordillera of southern Ecua-
dor and Peru.

RESUMEN

Al documentar la flora de la Reserva Natural Rio Nambi, se encontré eames especie,
Ardisia niambiensis. Se describe, se pee | pales y
Ardisia niambiensis se ubica mejor del subgé Ardisia debid icul inal
y gl. andular-papillos as con tamificaci cién tripinnada, su céliz simétticamente dividido con
lébulos simétricos y glandular-ciliolados a lo largo de los margenes. La nueva especie es
muy afin a Ardisia le tar una especie endémica a la falda oriental de la Cordillera
Oriental del Ecuador y Per

INTRODUCTION

The pantropical genus Ardisia Swartz contains approximately 400-500
species (Chen Cheih & Pipoly 1996) with centers of high species diversity
in Malesia and the Chocé Floristic Province from Panama through Co-
lombia to Ecuador (Pipoly 1991, 1992, 1994, 1995, 1996). Within the
Neotropics, approximately 300 species have been described, the majority

Stipa 18(2): 389-393. 1998

390 Sipa 18(2)

from Mesoamerica. There are relatively few South American taxa, perhaps
not passing 40 total. During the course of an analysis of the vegetation of
the Rio Nambi Natural Reserve, located in western portion of Narifio cor-
responding to the Chocé Floristic Province, a new species of Ardisia was
discovered, which is described herewith.

Ardisia niambiensis Pipoly & Cogollo, sp. nov. (Fig. 1). Type: COLOMBIA.
NARINO: Mpio. Barbacoas; Corregimientos Ortiz y Zamora; Vereda El Barro; Reserva
Natural Rio Nambi, ca. 5 dm al W de Al taquer, Faldas Occidentales de la Cordillera
Occidental, 01° 15’N, 78° 08’W, 1,250-1,350 m, 9 Sep 1997 (stam. Fl), /. Pipoly,
A. Cogollo, M. Lépez & M. Rodriguez 21572 (HOLOTYPE: PSO; ISOTYPES: BRIT, COL,
FMB, GH, HUA, JAUM, K, MO, NY, TULYV).

Ob inflorescentiam terminalem tripinnatipaniculatam, calycem aequaliter divisum, necnon
lobos calycinos simetricos glandul ari-ciliolaribusque subgeneri Ardisiae pertinet. Propter

oe on apice prev iacuminatas, rhachides inflorescentiares angulatos, margines sepalinos

premontano valde arcte affinis, sed ab ea ramulis
abs oe. fufuraceo- ae) Bet coriaceis (non chartaceis), subter glabris dense
punctatis punctato-lineatisque (nec furfuraceo-lepidotis), petiolis 7-10 (non 15-25) mm
longis, floribus pedicellatis (non sessilis), denique lobis calycinis deltatis vel late triangularibus
(non ovatis) 2—2.2 (nec 1-1.2) mm longis statim distinguitur.

Tree to 10 m X 15 cm DBH. Branchlets terete, 4-8 mm glabrous. Leaves
alternate; blades coriaceous, oblanceolate to obovate, (14.1—)19-—25.4 cm
long, (6.8-)7.5—9.4 cm wide, apically abruptly acuminate, the acumen 5—
8 mm long, basally acute, decurrent to petiole base, midrib canaliculate
and prominently raised above, prominently raised and densely black punctate-
lineate below, the secondary veins numerous, 39-4 pairs, prominulous
above and below, drying dark above, pallid and densely perpuncticulose
below, the punctations in the form of short lines, the margin slightly inrolled
except revolute basally; petioles marginate to base, 7-10 mm long, glabrous.
Inflorescence terminal, pyramidal, tripinnately paniculate, 12.8-22.5 cm long,
11.533 cm wide, peduncle 1—2.5 cm long, the rachis prominently angulate,
the secondary branches with smaller panicular units 12-16 cm long, 6-10
cm wide, the ultimate branches racemose, moderately rufous glandular-
granulose and prominently and densely black punctate-lineate; floral bracts
caducous, unknown; pedicels cylindrical, (1.0—)1.5—2.5(—3.0) mm long.
Flowers unknown, but fruiting calyx 3-3.5 mm long, rufous glandul papillate
and darkened at junction of lobe and tube, the lobes deltate to widely tri-
angular, 2-2.2 mm long, 2.2—2.5 mm wide, apex acute toa minutely acuminate
apex, minutely rufous-lepidote and carinate medially without, prominently
punctate and punctate-lineate, the margin hyaline, somewhat erose and sparsely
glandular-ciliolate apically. Fruit globose, 5—8(—10) mm long and wide when
dried; when fresh, exocarp fleshy, bitter tasting.

Distribution. —Known only from Reserva Natural Rio Nambi, Narifo,
Colombia, at 1,250—1,350 m elevation.

PIPOLY AND CoGoLLo P,, A new species of Ardisia 391

Fic. 1. Ardisia niambiensis Pipoly, showing the pyramidal tripinnate inflorescence, densely
and prominently black punctate leaf surfaces, short petioles marginate to base, and pedi-
cellate flowers. Drawn from the holotype.

392 Stipa 18(2)

Ecology—Ardisia niambiensis occurs in premontane pluvial forest, receiving
approximately 7,800 to 8,200 mm rain per year. The species occurs in relatively
low densities, in at the forest margin on ridgeto

Etymology,—Ardisia niambiensis is named phocnetiedly for the Reserva
Natural Rio Nambi, a private, nonprofit, integrated conservation and de-
velopment area in the state of Narifio, Colombia, operated by FELCA, the
Ecological Foundation of the Hummingbirds of Altaquer (Fundacion Ecoldégica
de los Colibris de Altaquer).

Paratype. COLOMBIA. Narino: Mpio. Barbacoas; Corregimientos Ortiz Zamora; Vereda
El Barro, Reserva Natural Rio Nambi, ca. 5 km al W de Altaquer, Faldas Occidentales de
la Cordillera Occidental, 01° 15’ N, 78° 08’ W; 1,350-1,450 m, 3 Sep 1997 (ster.), /.
Pipoly, A Cogollo, M. Lopez er M. Rodriguez 21227 (BRIT, FMB, JAUM, PSO)

On account of the shortly acuminate leaf apices, angulate inflorescence
rachises, carinate sepals with hyaline and subentire to erose margins, Ardisia
niambiensis is most closely related to Ardisia Nala Pipoly. It is sepa-
rated from A. premontana by its glabrous (not furfuraceous-lepidote) branchlets,
the coriaceous (not chartaceous) leaf blades that are densely black punctate
and punctate lineate (and not furfuraceous-lepidote) below, petioles 7-10
(not 15-25) mm long, pedicellate (not sessile) flowers and deltate or widely
triangular (not ovate) sepals that are 2.0—2.2 (and not 1.0—-1.2) mm long.

Ardisia subgenus Ardisia, to which A. niambiensis belongs, is defined by
the well-developed but early caducous floral and inflorescence bracts, race-
mose or rarely spicate branchlets of the inflorescence, the anthers dehisc-
ing by subapical pores opening into wide longitudinal slits, and symmetric
sepals. Within that subgenus, the other species known from the Chocé include
A. colombiana Lundell, A. granatensis Mez, A. opaca Lundell, A. valida Mez
(= A. perinsignis Lundell syn. nov.), A. monsalveae Pipoly, and A. cabrerae
Pipoly (Forero & Gentry 1990; Pipoly 1991, 1995) Among these species,
the only ones with inflorescence shape, pedicellate flowers and sepals simi-
lar to those of Ardisia niambtensis are A. cabrerae and A. monsalveae. How-
ever, the three species may be easily separated by the following key:

1. Calyx 2.4-2.9 mm long, apically obtuse, the margins glabrous, entire to
subentire.
2. Leaf blades coriaceous, elliptic; sepals 3.4-3.5 mm wide, glabrous ........ A. cabrerae
2. Leaf blades chartaceous, widely oblanceolate to obovate; sepals 0.5—0.6
mm wide, densely and minutely rufo-puberulent A. monsalveae
. Calyx 3-3.5 mm long, apically acute to a minutely acuminate apex, min-
utely rufous-lepidote and carinate, the margin sparsely ciliolate .......... A. niambiensis

—

ACKNOWLEDGMENTS

Support for our studies in plant diversity at the Rio Nambi Natural Reserve
is provided by a generous grant from the National Geographic Society, No.

PIPOLY AND CoGOL_o P., A new species of Ardisia 393

5575-95. We thank the collaboration of the herbarium of the Universidad
del Narifio (PSO), especially the Director, Bid]. Marta Sofia Gonzalez Insuasty,
and the herbarium technician, Bernardo Ramirez. We gratefully acknowl-
edge the logistical collaboration of FELCA (Fundacién Ecoldgica de los
Colibris de Altaquer), especially that of Don Marcial Bisbicuz and Mauricio
Flores. We also thank Juan Guillermo Ramirez Arango and Adriana Gomez
for technical and logistical help. We are most grateful to Dr. Ramiro Fonnegra,
director of HUA, for making the services of Gloria Mora available to us,
who skillfully prepared the illustration.
REFERENCES

CHEN, CuikH and J. Piroty. 1996. Myrsinaceae. Pp. 1-38. In: Wu Zheng Yi and P.H. Raven,
eds. Flora of China. Vol 15. Science Press, Beijing and Missouri Botanical Garden, St.
Louis, Missourt.

Forero G., E. and A.H. Gentry. 1989. Lista Anotada de las Plantas del Departamento
del Chocé, Colombia. Biblioteca José Jerénimo Triana, No. 10. Instituto de Ciencias
Naturales, Museo de Historia Natural, Bogota, D.E.

PiroLy, J. 1991. Notas sobre el género Ardisia Swartz (Myrsinaceae) en Colombia. Caldasia
16(78):277-284.

1992. Ardisia callejasii (Myrsinaceae): a new species from the Antioquian Chocé

ef Cslanibin Novon 2:389-391.

. 1994. New ces oie (Myrsinaceae) from the Cordillera Occidental of Colombia

and Ecuador. Novor 4,

. 1995. Dos nuevas especies del género Ardisia (Myrsinaceae) de la provincia floristica
chocoana de Colombia. Caldasia 17(82—85):419-424.

. 1996. New species of Ardisia (Myrsinaceae) from Ecuador and Peru. Sida 17:445—
458.

ie

394 Sipa 18(2)
BOOK REVIEW

THOMAS LUMPKIN and Dean McC ary. 1994, Azuki Bean: Botany, Pro-
duction and Uses. (ISBN 0 85918 765 6, hbk). CAB International,
Oxford University Press, 198 Madison Ave., New York, NY 10016.
800-451-7556. $70.00. 268 Pp.

This book purports to be a monograph, including both the basic botanical and applied
agr onomic aspects among the “Azuki” members of Vigna subgenus Ceratotropis. The book
is divided into nine chapters, ane: an introduction, botany, physiological character-
istics, production, j insects and nematodes, diseases, breeding characteristics, food chem-
istry and processing, and uses a marketing. The agronomic chapters contain a thor-
ough review of the literature, but much of it refers only to the Azuki beans in general, and
not to a specific taxon. As a plant systematist, | am most disappointed with Chapter 2,
the Botany of Azuki. | find it superficial, and not useful for understanding the differences
gues the species, or the infraspecific taxa (subspecies, varieties and cultivars). A synop-

s or taxonomic revision for this relatively small group could have been saree by
any one of a number of qualified taxonomists, but the authors chose to avoid a formal
treatment. They state that in their Table 2.1, “No attempt has been a to correct the
confusion of different classifications in the literature.” That statement amounts to a fatal
flaw for the work, because many of the subsequent chapters make frequent reference to
various cultivar names, without link to a taxon described in the Botany chapter. For ex-
ample, in the discussion of starch ¢ granules and processing (Chapter 8), reference is made
to “azuki cv Takara,” be there is no mention of that or any other cultivar in the taxonomy
chapter. Because the reader is most often unsure of the exact entity referred to, the rest of
the book loses much of i its usefulness, except in broad terms. I find the book hardly more
than a literature review, and not a definitive monograph. Its extensive ee may

be useful for agronomic investigators, but hardly justifies its hefty price. | would have
preferred to ee seen a carpieas po executed — sacle omic and agri-
cultural treatment, like the classic work of L. van der 1 for Cicer arietinum, pub-

lished in Meded. Landbouwhogeschool ae 10:1-342. 972. | highly recommend that
Maesen’s monograph be used as a model (with ene systematics added) for eco-
nomically critical and highly domesticated plant taxa. By following Maesen’s model, we
may better take advantage of wild relatives of currently cultivated plants, to bring useful
characteristics into the cultivated from the wild. The book a be part of any major
agricultural library, as a literature review source.—John Pipoly, IT

SDA 18(2): 394. 1998

CLUSIA NIAMBIENSIS (CLUSIACEAE),
A NEW SPECIES FROM THE CHOCO FLORISTIC
PROVINCE OF COLOMBIA AND ECUADOR

JOHN J. PIPOLY UI

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
Jpipoly@brit.org; clusia@latinmail.com

ALVARO COGOLLO P.

Fundacion Jardin Botinico, Joaquin Antonio Uribe
Apartado Aéreo 21407, Medellin, Antioquia, COLOMBIA
acogollo ‘L.com; jardinbo @epm. net.co
MARTA SOFIA GONZALEZ

Departamento de Biologia, Universidad de Narifio
Apartado Aéreo 1176, Paste, Narino, COLOMBIA

ABSTRACT

Documentation of the flora of the Natural Reserve Rio Nambf has resulted in the dis-
covery of a heretofore undescribed species, Clusia niambiensis. The species is described,
illustrated and its salient morphological features are elucidated. While its vegetative mor-
phology is reminiscent of both Clusia laurifolia and C. venusta, the free, numerous, and
subulate stamens with a central mass of staminodia producing a waxy resin indicate it is a
member of section Chlamydoclusia rather than section Retinostemon, where the aforemen-
tioned congeners belong. At present, the relationships of Clusia niambiensis within sec-
tion Chlamydoclusia, are unknown. It is anticipated that continuing studies of the genus,
leading to a treatment of the family for Flora de Colombia, will resolve this problem.

RESUMEN

ie

cumentar la flora de la ae Neate Rio Nambi, se encontré una nueva especie,

Clusia niambiensi 1S Se describe obresaliente

Mientras que sus caracteres vegetativos son muy semejantes a los de Cues laurifolia y C.
venusta, sus estambres libres, numerosos y subulados indican claramente que C. niambiensis
ase a la seccién Dee aes y no a la seccién Eat OTh donde se ubican las

la ubicacién
taxonémica de Clusia niiambiensi ee de la seccién ere ne Esperamos que a
través de los estudios en proceso para el tratamiento taxonémico de la familia para Flora
de Colombia, se pueda resolver este problema.

Recent exploration of the Reserva Natural Rfo Nambj, located on the

SIDA 18(2): 395-399. 1998

396 Sipa 18(2)

western slopes of the Western Cordillera of the Colombian Andes, has re-

sulted in the discovery of this spectacular new species, described herewith.

Clusia (§ Chlamydoclusia) niambiensis, Pipoly, Cogollo et Gonzalez, sp.
nov. (Fig. 1). TyPE: COLOMBIA. NARINO: Mpio. Barbacoas; Corregimientos Ortiz
y Zamora; Vereda El Barro; Reserva Natural Rio Nambi, ca. 5 km al W de Altaquer,
Faldas Occidentales de la Cordillera Occidental, 01° 15' N, 78° 08' W; 1,450-1,500
m, 8 Sep 1997 (stam. Fl), J. Pipoly, A. Cogollo, M. Lopez & M. Rodriguez 21489 (HO-
LOTYPE: PSO; ISOTYPES: BRIT, COL, FMB, JAUM, K. MO

Quoad lamina pure nesvatiosque EsOe vas Ones resinam alba a waked ae
nia

similans sed ab ea petiolis f

—

ad bases laminas acutas decurrentesque (non sine he sad eae nas tia fr Gaia
ovoideis vel ellipsoideis (non subglobosis) praeclare distat. Propter stamina inter se libera
subulataque centraliter resinifera, sn ata conniv entaque sectio Chlamydoclusia pertinet,

inter species aliis petiolis prof -ulatis arginatisque, laminis bullatis nervatiosque

brochidodromis, Gaiceibae longitudinaliter costatis statim distinguitur

Glabrous, dioecious, shrubs or treelets to 2—3(—5) m tall, aout dynam-
ics corresponding to Scarrone’s Model; latex white. branchlets terete, ap-
pearing tetragonal when dried, 8—-10(—15) mm diam., the bark reddish-
brown, exfoliating transversely in the upper nodes, appearing furfuraceous
at times. Leaves decussate; blades chartaceous, obovate, (19—)27—38.5 cm
long, (14.5—)15.2—23 cm wide, apically rounded to obtuse, basally acute,
decurrent to the petiole base, bullate, the midrib prominently raised or raised
within a depression above, prominently raised below, the secondary veins
brochidodromous, 28-46 pairs, ca. 7-10 mm apart, alternating so every
other vein has twice the diameter of the others, deeply impressed above,
prominently raised below, the submarginal connecting vein ca. 2-4 mm
from margin, glabrous above and below, at times when dried with small
linear latex canals oxidizing red, the margin scarious, entire; petioles deeply
canaliculate and marginate, (1.0—)2.5—3.5 cm long, without adaxial mar-
gined pit. Staminate inflorescence erect in bud, pendulous at maturity, a compound
cyme, 20-25 cm long, 10-20 cm wide; peduncle appearing tetragonal when
dried, 10.5—17 cm long, the bark transversely checked and exfoliating; primary
inflorescence bract coriaceous, lanceolate, 18-21 mm long, 4-6 mm wide,
apically acute, the midrib prominently raised above and below, the margin
scarious, entire, glabrous; secondary inflorescence bracts cartilaginous, sub-
orbicular, 3.5—4.0 mm long, 3.8—4.2 mm wide, apically broadly rounded,
medially carinate, the margin scarious, entire; tertiary and floral bracts as
in secondary but smaller acropetally, to the smallest that are rhomboid, 3
mm long and wide, apically acute; bracteoles 2, cartilaginous, ovate, .3.8—
4.0 mm long, 2.5—3.0 mm wide, apically rounded, cucullate, the margin
hyaline, not scarious, entire; pedicels 1—2.5(—5) mm long. Staminate flow-
ers sepals 8, the lower 4 opposite, the upper 4 contorted, coriaceous, acro-

PIPOLY AND CoGoLLo P., A new species of Clusia BOF

1 af

oy aX
LQ

Fic. 1. Clusia niambiensis Pipoly, Cogollo & are A. Els B. Sjereies! pistillate Hes
showing staminodes and stigmatic area. C. Opened

central eater mass. D. Fruit, acts costae and mature stigmas. Figs. A-D, aes
from holotype

398 Sipa 18(2)

petally larger, the lowermost oblate, 4.4-5.5 mm long, 5.5—6.5 mm wide,
apically broadly rounded, translucent toward the margins, the linear latex
canals black, conspicuous, the margins flat, scarious, irregularly notched,
at times also appearing erose; uppermost sepals as in lowermost but ob-
long, 10-12 mm long, 7.5—8 mm wide, petals 12-16, contorted, mem-
branaceous, oblong to oblanceolate, acropetally larger, the largest 16-19
mm long, 10-12 mm wide, somewhat clawed, some appearing oblanceolate-
spathulate, apically broadly rounded to truncate, linear resin canals con-
spicuous, black, numerous, the margin undulate, somewhat irregular, mi-
nutely erose at tip; stamens numerous, linear 4-5 mm long, the filaments
ae 2.0—2. ; mm long, the anther sacs linear, 2.0-3.0 mm long, the con-
0.5 mm long, subulate; central staminodes
forming a mass ; agplutinated by orange waxy resin, the pistillode vestigial
or absent. Pistillate inflorescence as in staminate but peduncle 7—20 cm long;
secondary inflorescence bracts 11—13 mm long, 5.5—6.6 mm wide in lower
portion of inflorescence, acropetally smaller to 3 mm long, 3 mm wide;
pedicels accrescent to 2.8 cm in fruit. Pistillate flowers as in staminate but
staminodes 4—5 mm long, devoid of pollen, the connectives 0.6-1 mm long,
pistil ovoid, 7-8 mm long, 8-9 mm wide, carpels and stigmas 8, cunei-
form at maturity, convex, 4.5—5.2 mm long, 2.0-2.5 mm wide. Fruit ovoid
to oblong, 3.0-6.5 cm long, (1.8—)2.7—2.9 cm diam., 7-ribbed, the fruit
grenadine, the ribs white when fresh.

Distribution.—Endemic to the Chocé Floristic Province of Colombia
and adjacent Ecuador, (17—)650—1,650 m elevation.

Ecology and conservation status. —Clusia niambiensis occurs in premontane
and montane pluvial forest, where it is an understory treelet. The popula-
tions of this species are composed of approximately 20 individuals per hectare.

Etymology— Clusia niambiensis is named for the Reserva Natural Rio Nambi,
a private Wildlife Reserve in the state of Narifio, Colombia, operated by
FELCA, the Ecological Foundation of the Hummingbirds of Altaquer, where
the largest populations of this species have been found.

Local names and uses.—*Guandera” (Colombia & Ecuador, Spanish).

ParatyPes: COLOMBIA. NariNo: Mpio. Barbacoas; Corregimientos Ortiz y Zamora;
Vereda El Barro; Reserva Natural Rio Nambi, ca. - km al W es Altaquer, Faldas Occidentales

de la Cordillera Occidental, 01° 15' N, 78° 08' W; 1,450-1,500 m, 1 Sep 1997 (pist. fl),
J. Pipoly, A. ea et al. 21044 (BRIT, va oe 8 Sep ee ae fl), J. Pipoly, A
Cogollo, M. Lopez & M. Rodriguez 21490 (BRIT, COL, JAUM O), 1,250—1,350 m,

10 Sep 1997 (stam. fl), J. Pipoly, A. Cogollo et al. 21655 (BRIT, ov PSO); Resguardo
de El Sabalo, Rio Cangapi, 01° 17' N, 78° 14' W, 580 m, 17 Aug 1995 (fr), B. Ramirez et
al. 8049 (BRIT, PSO); orceimicnic i Pee, near e | Diviso, 25 Nov 1979 (piste. fl),
O. de Benavides 2149 (PS O); Corregimiento de Jun 200 m, 7 Oct 1988 (pist. fl bud),
O. de Benavides 10246 (PSO); Locality lees Junin a Divisio, 1,700 m, 15 Sep 1978

Pipoty AND CoGoLto P., A new species of Clusia 399
(pist. fl), O. de Benavides 1565 (PSO); Espino to Tumaco Road, 84 km W of Espino, on
pass between Altaquer and Junin, 01° 15' N, 78° 09’ W, 1,300 m, 18 Nov 1986 (fr), B
Hammel & R. Bernal. 15751 (COL, HUA, MO, PSO); El Espino to Tumaco Road, 30
km W of Ricaurte, 10 km W of Altaquer, El Mirador, Finca Sta. Lucia, in forest N of Rio
Nambi, 01° 17' N, 78° 07' W, 950 m, B. Hammel & A. Narvdez 17178 (MO, PSO); Mpio.
Ricaurte, Hacienda La Planada, 1,850 m, 26 Nov 1981 (fr), O. de Benavides 3338 (PSO);
Resguardo Indigena Nulpe Medio, Andalucia- Nulpe Medio, 01° 05' N, 78° 14’ W 01°
18'N, 77° 54' W, 800-1,100 m, 5 Jan 1996 (pist. fl), B. Ramirez et al. 9364 (BRIT, PSO);
E] Diviso, highway to Tumaco, 790 m, 20 Feb 1968 (fr), G. Lépez 261 (PSO); Mpio. de
Tumaco, 2 km from Tangareal, banks of Rio Mira, 17 m, 8 Apr 1978 (pist. fl), O. de Benavides
1383 (PSO). ECUADOR. Carcut: Cantén Tulcaén, Reserva Indigena Awa, Gualpi Alto
Community, Parroquia Chical, 01° 02' N, 78° 14' W, 1,800 m, 15-28 Jul 1991 (stam. fl
bud), D. Rubio et al. 1589 (BRIT, MO, QCNE), (fr), D. Rubio et al. 1590 (BRIT, MO,
QCNE). Los Rios: Cantén Quevedo, Parroquia Centinela-La Pirdmide, via Sto. Dominto
de los Colorados-Quevedo, entrando por Patricia Pilar, km 41, 01° 40'S, 79° 20' W, 650
m, 25 Feb 1992 (fr), C. Quelal & G. Tipaz 160 (BRIT, MO, QCNE). PICHINCHA: Quito-
Puerto Quito Road, 10 km N of main road, Km 113, 00° 05' N, 79° 02' W, Reserva ae
estal ENDESA, Rio Silancha, Cooporacién Forestal Juan Manuel Durini, voucher for NC
650-700 m, 17 May 1987 (pist. fl), P Acevedo R., D. Daly & M. Rios 1695 (BRIT,
QCA, US).

Clusia niambiensis is infrequent in the western slopes of the Cordillera
Occidental of Colombia and adjacent Ecuador, where it has been frequently
confused with its congeners, Clusia laurifolia P\. & Tr., and Clusia venusta
Little, all of them having bullate leaves with obviously brochidodromous
venation. However, the free stamens with subulate apices, and resiniferous
zone in the center of the androecium indicate it is a member of section
Chlamydoclusia, and not section Ketinostemon, where the other two taxa
belong. At this time, it is not known which member of subgenus Clusia is
its closest relative.

ACKNOWLEDGMENTS

This work is a result of a generous grant from the National Geographic
Society, 5575-95, for exploration of the Reserva Natural Rio Nambi. We
thank Bernardo Ramirez (PSO) for his technical assistance, as well as the
administration of FELCA (Fundacién Ecoldgica de los Colibris de Altaquer),
Marcial Bisbicuz, and Maruricio Flores. Ramiro Fonnegra, Director of HUA,
graciously made the services of Gloria Mora available to us to prepare the
line illustration, for which give thanks.

400 Stipa 18(2)
BOOK REVIEW

CHRISTINE PADOCH and Nancy Ler PrELuso, eds. 1996. Borneo in Transi-
tion: People, Forests, Conservation and Development. (ISBN 967-
65-3110-3 (hbk). Oxford University Press, 198 Madison Ave., New
York, NY 10016. 800-451-7556. $55.00. 291 pp, tables, figures, maps.

This book is a collection of papers resulting from a symposium “Interactions of Peop dle

and Forests in Kalimantan” held at the New Yc ae Laine Garden it 11991. Despite be-
ost

ing published some five years after the event, the t {therein have not
their timeliness, and are synthesized by an cine: pile chapter by Padoch and

Peluso. The book is composed of three sections: 1) Introduction to Conservation and Development
and the Role of Research, I) Local Transformations in Forest Use and Forest Rights,
and III) Case-Studies in Resource Management

The first section contains five papers that explain the current development of Borneo
within the context of Indonesia, Malesia as a whole, and SE Asia. Of these, the chapter by
Potter on forest degradation is particularly illuminating and logically outlines what the
conservation situation was on the island (prior to El Nifo-1998), although the debt for
nature swap, sensu Potter, is an extremely controversial concept.

The second section is focused to development on a local level and its effects on the for-
est, its people, and their sociology. It was interesting to note that Map 9.2 (p. 140) in Ngo’s
article, showing the migratory direction of the Kayan i in Kalimantan, lies eae the conti-
nental divide (and the Sarawak Border) and is immediately adjacent to (east of) and slightly
re with, the area surveyed by Burley in Chapter 5. Chapter 8, by Peluso and

Padoch, is a cogent analysis of how two Dyak villages have changed with development,
and contains a new hypothesis sree changes j in “distribution and economic impor-

tance of particular crops and technologie
and rights. It includes extensive ane ee a Pe with a eqncasune section
that Sea the areas where further research is needec

re third part, Peters (Chapter 14), in his article on Ilipe Nuts, has provided a model
case ne I hope a rapidly adopted standard protocol) for the study of underexploited tropical
crops. If data similar to that presented by Peters are combined with market data using the
methodology in the following Chapter 14, by Leaman et al., a realistic estimation of a
crop’s potential can be calculated. Understanding the biology, and therefore, the likeli-
hood of cultivation success for a given crop, combined with its relative “ svi fit-
ness” against other crops siaiilianeotsly available in the market, bins be a desirable for-

—_

mula to guide local communities in preferential resource allocatior

In summary, this book provides not only information Sones to cote: but also contains
papers with protocols worthy of becoming standards for their respective fi elds, concomi-
tant with rich bibliographies. I think every conservationist, economic botanist, sustain-
able development specialist, and resource manager should have a copy.—/ohn J. Pipoly III

Stipa 18(2): 400. 1998

NEW SPECIES AND NOMENCLATURAL NOTES
IN CLUSIA (CLUSIACEAE) FROM ANDEAN
COLOMBIA AND VENEZUELA

JOHN J. PIPOLY II

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76126-3019, U.S.A.
Jpipoly@brit.org; clusia@latinmail.com

ALVARO COGOLLO P.

Fundacion Jardin Botdnico, Joaquin Antonio Uribe
Apartado Aéreo 51407, Medellin, Antioquia, COLOMBIA

acogollo @lati l.com; jardinbo @epm.net.co

ABSTRACT

Continuing studies on the Clusiaceae for Flora de Colombia, and fieldwork to docu-
ment the phytodiversity of Rio Nambi Natural Reserve have revealed two new species,
Clusia bernardoi Pipoly & Cogollo, and C. detragona Pipoly & Cogollo. a8 ae new species
are described, illustrated, and hypotheses of their f J. Corrections
to bibliographic errors in citations for Clusia sect. Oede -matopus and Clusia sect. Havetiopsis
are provided. The new combination Clusia comans (Mart.) Pipoly is validated. Colombian
and related Andean Venezuelan species formerly included in the genus Oedematopus are
transferred to Clusia, necessitating the new combinations: C. aristeguietae (Maguire) Pipoly,
C. divaricata (Cuatrec.) Pipoly, C. epiphytica (Cuatrec.) Pipoly, C. mirandensis (Cuatrec.)
Pipoly.

RESUMEN

Estudios hacia un tratamiento taxonémico de la familia Clusiaceae para Flora de Co-
lombia y para documentar la fitodivdersidad de la Reserva Natural Rio Nambi, revelaron
os especies nuevas para la ciencia, Clusia bernardoi Pipoly & Cogollo, y C. tetragona Pipoly
cee ogollo. Se describen, se ilustran y se proponen hipdoteses de parentezco para las nuevas
especies. Errores bibliograficos previamente publicados para Clusia secciones Oedematopus
y Ha 5 ee La poe combinacién ae comans (Martt.) Bey . asa
la regién Andin

miembros del género Oe se transfieren a Clusia, saree en las nuevas ca aa
C. aristeguietae (Maguire) Pipoly, C. divaricata (Cuatrec.) Pipoly, C. epiphytica (Cuatrec.)
Pipoly, C. mirandensis (Cuatrec.) Pipoly.

INTRODUCTION

The genus Clusia contains approximately 300 species, of which nearly
17% remain undescribed. Fundamental taxonomic work by Panchon and

SiDA 18(2): 401-409. 1998

402 Stipa 18(2)
Triana (1860a, 1860b), Engler (1888, 1895), and Vesque (1892, 1893) form

the basic framework which all modern studies have been based to date.
Cuatrecasas (1949, 1950) was the first author of this century to critically
reevaluate the Colombian members of the family, and in the two afore-
mentioned papers, he described a total of 75 taxa. Most of the taxa de-
scribed by Cuatrecasas belonged to the genus Clusia sensu stricto, but his
work also included critical additions to the genera Oedematopus and Clusiella
that left the circumscription of each significantly emended. Subsequently,
Ewan (1951), Maguire (1951, 1977), Pipoly and Graff (1995a, 1995b),
Pipoly (1997) and Pipoly and Graff (1995a, 1995b), described a number
of new species. Pipoly et al. (1998), in their treatment of the family for
Flora of the Venezuelan Guayana, relegated Quapoya, Havetiopsis, and Oedematopus
(among other genera) to synonymy under C/usia. This paper is intended
to describe novelties in Clusia, correct bibliographic errors in citation for
sections Havetiopsis and Oedematopus, validate the new combination Clusia
comans, and to complete the transferal of Oedematopus species to Clusia.
NOVELTIES IN CLUSIA
In the course of identification of herbarium materials for an analysis of
the family for Flora de Colombia, the following new species were encoun-
tered.
Clusia (§ Anandrogyne) bernardoi Pipoly & Cogollo, sp. nov. (Fig. 1).
COLOMBIA. CAuCa: Mpio.de Tambo, Parque Nacional Munchique, km 60-80 via

ala Galera, 1,850—2,150 m, 13 Apr 1995 eel fl), B. Ramirez, N. Rojas, L. Zambrano,
N. Diago 7198 (HOLOTYPE: PSO; ISOTYPES: CAUP, JAUM)

Quoad filamenta numerosa omatne liber: aad apices ome argine at, a aGaNG ba ases See ata,

androphoros Ps auciprae edita C. tio rg’) I I
oe Jo : q .

vel peranguste oblanceolatis, |
] : 15 1:

is vel late rorundatls
hydropoditsque indutis jue 4 perfacil .

palis | p 8
Glabrous treelet to 5 m - latex unknown. Branchlets terete, 5-8 mm
diam., swollen at the nodes 7-10 mm diam., the nodes short, 1.3—2 cm
long, glabrous. Leaves sessile; blades coriaceous, oblong to narrowly oblan-
ceolate, 17-23 cm long, 4.6-6.0 cm wide, apically truncate to broadly rounded,
basally cuneate, midrib prominently raised above and below, the second-
ary veins brochidodromous, numerous, 70—80 pairs, prominulous above,
— visible below, the submarginal collecting vein like the secondaries,
mm from margin, drying smooth and dark green above, pallid below,

om linear resin canals visible in the lower 1/3, and with scattered, minute
rubiginous hydropotes, the margin slightly revolute except prominently revolute
in the basal 1/4 of leaf. Staminate inflorescence a terminal panicle of cymes,
thrice-branched, and each subsequent branch divided 3 times; peduncle
3.74.7 cm long; inflorescence bracts cartilaginous, ovate, 7-10 mm long,

PIPOLY AND COGOLLO P., New species and nomenclatural notes in Clusia 403

Imm
Fic. 1. ae bernardoi od tase A. Branchlet, showing obl
leaf blades. B. Androecium, showing poorly developed androphore and hea Were
with ue apices. C. Sepal, showing asymmetry, and conspicuous venation. D. Petal,

showing asymmetry and conspicuous venation. E. Abaxial ene showing hydropote.

A-—D, drawn from holotype.

404 Sipa 18(2)

4.5-6.5 mm wide, apically obtuse, medially thickened but not carinate,
the linear resin canals drying black, conspicuous, the margin scarious, en-
tire; floral bracts 2, as in secondary inflorescence branch bracts except ob-
late, 1.8-2.2 mm long, 2.1—4.1 mm wide, apex widely rounded; bracteoles
2, chartaceous, oblate to deltate, 2.0—2.2 mm long, 3.8-4.2 mm wide, apex
obtuse to acutish, medially keeled, the margin entire; pedicels obsolete. Staminate
flowers 3 per cymule, whitish-green; sepals 4, decussate, membranaceous,
oblong, 5.5—6.5 mm long, 3.1-3.5 mm wide, asymmetric, apically obtuse,
cucullate, hyaline, the venation conspicuous, the margin entire; petals
decussate, oblong, 5.5—7.0 mm long, apically broadly rounded and asym-
metrically notched, hyaline, the venation conspicuous, the margin irregu-
lar, entire; androphore poorly developed or obsolete, stamens 22-28, 2.0—
3.5 mm long, free to the base, the filaments 1-1.5 mm long, the anthers
oblongoid, 0.5—0.9 mm long, 0.5—0.7 mm wide, apically emarginate, ba-
sally subcordate, as wide as the filaments, deshiscent by wide longitudinal
slits throughout their length; pistillode absent. Pistillate inflorescence, flow-
ers, and fruit unknown.

Distribution.—Known only from the type.

Ecology and conservation status.—The type locality lies in a transitional
zone between premontane pluvial forest and cloud forest, but no further
details are known. While the species is known from Munchique National
Park, encroachment is common and poses a threat to this species.

Etymology.—It gives me great pleasure to dedicate this striking new spe-
cies to Bidl. Bernardo Ramiro Ramirez Padilla, Herbarium Technician of
the University of Narifio in Pasto, Colombia. Bernardo has worked inde-
fatigably for over twenty years, documenting the complex, species-rich and
biogeographically important flora of the Department of Narifio and adja-
cent areas, such as Cauca and Putumayo.

Clusia bernardoi is unique among members of C. section Anandrogyne
because of its oblong to narrowly siemeeetae leaf blades bearing rubigi-
nous hydropotes, and 4-merous perianth. Four-merous flowers are other-
wise frequent among members of C. section Criuva (Pipoly, 1996), but the
oblongoid anthers with emarginate apices and cordate bases, dehiscent by
wide longitudinal slits leave no doubt that Clusia bernardoi is a member of
C. section Anandrogyne, the largest and most complicated section of the
genus.

Clusia (§ Criuva) tetragona Pipoly & Cogollo, sp. nov. (Fig. 2). COLOMBIA.
NARINO: Mpio. Barbacoas; oe mane year i auanaay Reserva
Natural Rio Nambf; ca. 5 km W de Altaquer, fal ; de la Cordillera Oc-
cidental; 01° 18' N, 78° 08' W, 1,350 ne 400 m, 3 Sep 1997 (bud, fr), /. Pipoly, A
Cogollo, M. Lopez, & M. Rodbiouee 21198 (HOLOTYPE: PSO; ISOTYPES: BRIT, CO L,
FMB, JAUM, kK, MO, NY, TULY).

PIpPOLy AND COGOLLO P., New species and nomenclatural notes in Clusia 405

Fic. 2. Clusia tetragona Pipoly & Cogollo. A. Branchlet, showing tetragonal shape appear-
. . 5 . ~

ing alate, and subsessile leaves. B. Androecium, showing poorly developed androphore,
free filaments, and muticous anthers. C. Flower buds, showing secondary inflorescence
bracts and two of the four bracteoles. D. Fruit, showing rostrate apex. A-C, drawn from

holotype. D, drawn from /. Pipoly et al. 21157.

406 Sipa 18(2)

Propter lamina obovata vel ase paket nervio submarginale prominente,
inflorescentiam terminalem pyramido-panicula ructu rostrato
C. garciabarrigae valde arcte affinis, ed ab ea ramulis tetragonis (non teretibus), petio
obsoletis vel usque ad 1.5 (non 2.5-4) cm longis, bracteis florinis 4(non 2), aie
(non oblongis), denique fructibus 8—10(—13) (non 5—7) mm longis statim distinguitur.

_

Terrestrial tree to 25 m tall, and to 45 cm DBH; latex white. Branchlets
tetragonal, subalate, (4—)5—-7 mm diam., glabrous. Leaves decussate; blades
coriaceous, obovate to elliptic, (9.5—)14.2-19 cm long, 4.5—G6(—9.5) cm wide,
apically broadly rounded to truncate, basally acute, midrib prominently
raised above and below, the secondary veins numerous, 50—52 pairs, con-
nected by a submarginal collecting vein ca. 2 mm from margin; linear latex
canals not visible from above, rubiginous and conspicuous below, the mar-
gin entire, flat; petioles obsolete to broadly marginate, to 1(—1.5) cm long.
Staminate inflorescence a bipinnate panicle, ca. 12 cm long, 10 cm wide;
peduncle 4—4.5 cm long, tetragonal except drying flat in the upper por-
tion; primary inflorescence bracts 2, cartilaginous, 11-13 mm long, 5.5—
6.5 mm wide, apically obtuse, medially keeled, the margin scarious, entire;
secondary inflorescence bracts acropetally smaller and similar, except the
uppermost suborbicular, ca. 3 mm long and wide; pedicels obsolete to 0.5
mm long. Staminate flowers greenish white; bracteoles 4, cartilaginous, oblate,
2.5-3 mm long, 3—3.5 mm wide, apically acute to rarely subobtuse, prominently
carinate medially, the margin entire, opaque, glabrous; sepals 4, membra-
nous, hyaline, decussate, the outer suborbicular, 4.5-5.5 mm long, 3.5—4
mm wide, the inner oblong, 7-8 mm long, 3.5—4 mm wide, apically cu-
cullate and broadly rounded, the linear latex canals conspicuous, the mar-
gin entire; petals 4, decussate, membranaceous, obovate-spathulate, 8—9
mm long, 4-5 mm wide, linear latex canals conspicuous, the margin en-
tire; androphore poorly developed, to 1 mm high; stamens 32-36, 3—4.5
mm long, the filaments free, angulate 1-2 mm long, the anthers linear, each
theca half the diameter of the connective, apically muticous, tapering ba-
sally into filament, dehiscent by narrow longitudinal slits 2-2.5 mm long;
pistillode absent. Pistillate flowers similar to staminate except somewhat smaller
(in bud); staminodes obsolete; pistil oblongoid, 3.5—4 mm long, ca. 2 mm
wide, the stigmas 4, sessile, cuneiform, concave, 0.9-1.3 mm long, 0.9-
1.1 mm wide, appearing to be borne on connivent styles at maturity, the
apparent styles 1-1.2 mm long. Fruit ellipsoid, 8—10(—13) mm long, 4.5—
6 mm wide, abruptly constricted to a small “beak” apically (rostrate).

Distribution.—Apparently endemic to the Barbacoas area, on the west-
ern slopes of the Cordillera Occidental, in the Department of Narifio, Colombia,
at 1,325—-1,900 m elevation.

Ecology and conservation status. —Clusia tetragona is restricted to premontane
pluvial and cloud forests, where it is found along forest margins.

Etymology.—The specific epithet, ‘tetragona’ refers to the tetragonal na-

PIPOLY AND CoGOLLo P.,, New species and nomenclatural notes in Clusia 407

ture of the branchlets when seen in transverse section.
Common names.—*Guandera” (Spanish).

ParatyPes. COLOMBIA. Narino: Mpio. Barbacoas, Cee Altaquer, Nis
El Barro, Reserva Natural Rfo Nambi, left bank of Rio Nambi, 01° 18'
1,325 m, 11 Dec 1993 (fr), PB Franco et al. 5145 (COL, PSO); Corregimiento a iz :
Zamora; Vereda El Barro; Reserva Natural Rio Nambf; ca. 5 km W de Altaquer, faldas
occidentales de la Cordillera Occidental; 01° 18' N, 78° 08' W, 1,350-1,400 m, 2 Sep
1997 (bud, fr), /, Pipoly, A. Cogollo, M. Lépez & M. Rodriguez 21157 (BRIT, COL, FMB,
JAUM, K, MO, PSO, TULV); Mpio. Ricaurte, La Planada, 1,900 m, 28 Nov 76 (stam.
fl), O. de Benavides 730 (PSO), La Planada, 5 km S of Altaquer, 01° 10’ N, 78° 00' W,
1,750 m, 22 Nov 1986 (fr), B. Reig as Bernal 15802 (COL, MO, PSO); La Planada,
7 kon de Chucunés; 01°:10°N; 77° 58’ W,.1,800 mm; 25 Sep 1989 Gr),O de Bee
10803 (MO, PSO), 18 Jan 1990 (fr), O. de Benavides 11337 (MO, PSO), 01° 05'N
01' W, 1,800 m, 22 Dec 1987 (fr), A. Gentry & P Keating 59721 (MO, PSO); Tail to i
Hond6n, 5—12 km SW of La Planada, 01° 04' N, 78° 02' W, 1,750—1,800 m, 6 Jan 1988
(fr), A. Gentry & P Keating 60474 (MO, PSO); Camino Las Cruces-Curucel, 01° 08' N,
77° 51' W, 1,700-1,800 m, 5 Nov 1995 (fr), B. Ramirez, M. Gonzdlez e& A. Munoz 8692
(BRIT, PSO).

Because of its obovate to elliptic, coriaceous leaves, with a prominent
submarginal collecting vein, terminal pyramidal-paniculate inflorescence,
and rostrate fruit, Clusia tetragona is most closely related to C. garciabarrigae
Cuatrecasas. However, Clusia tetragona is immediately separated from that
species by the tetragonal branches, appearing alate when dried, sessile to
short-petiolate leaves, more numerous floral bracteoles, suborbiculate se-
pals and much longer fruits. While Clusia tetragona is apparently sympat-
ric with C. garciabarrigae, it is notable that C. tetragona is restricted to more
open habitats than C. garciabarrigae. Clearly, more fieldwork is needed to
understand the population biology of these species.

NOTES ON CLUSIA SECTIONS OEDEMATOPUS AND HAVETIOPSIS

In our recent treatment of this section for the Venezuelan Guayana (Pipoly
et al. 1998), we inadvertently missed the first publication of the basionyms,
Oecedematopus Planch. & Triana, and Havetiopsis Planch. & Triana, and cited
them as “Ann. Sci. Nat., Bot. ser. 4, 14:249. 1860,” for Oedematopus and
“Ann. Sci. Nat., Bot. ser. 4, 14:246. 1860” for Havetiopsis, where complete
descriptions of the genera were presented, followed by descriptions of all
infrageneric taxa. However, Planchon and Triana had first published the
two generic names, with diagnoses inside a synoptic key, for the first time
in the previous volume, published in the same year. For my relegation of
Pilosperma Planch. & Triana to synonymy under Clusia, | (Pipoly 1997)
cited “Ann. Sci. Nat., Bot ser. 4, 13:315. 1860,” which was correct. Be-
cause the citations for Oedematopus and Havetiopsis only involve bibliographic
errors (ICBN, Art. 33.3), the combinations are valid, but should be cited

as follows:

408 Sipa 18(2)

= L. sect. Oedematopus (Planch. eee Pipoly in Steyermark, Berry
Holst, Fl. Venez. Guayana 4:269. 1998. Ocedematopus Planch, & Triana,
ee Sci. Nat., Bot. ser. 4, 13:315. 1860. one Havetia octandra Poepp. & Endl.,
Nov. Gen. Sp. 3:11. t. 209A. 1840 (LEC ao ib — -ipoly 1998). = Clusia octandra
(Poepp.) Pipoly, Fl. Venez. Guayana 4:276.

Clusia L. sect. Havetiopsis (Planch. & Triana) Pipoly in Steyermark, Berry
& Holst, Fl. Venez. Guayana 4:269. 1998. Havetiopsis Planch. & Triana,
Ann. Sci. Nat., Bot. ser. 4, 13:315. 1860. LECTOTYPE SPECIES, here designated: Clusia
flavida (Benth.) Pipoly, based on Havetia flavida Benth., London J. Bot. 2:369. 1843.

When relegating Renggeria to synonymy under Clusia, we (Pipoly et al.
1998), cited the basionym “Renggeria comans Meisn., Pl. Vasc. Gen.,

Commentarius 42. 1837.” However, Meisner did not make a new doen

nation on that page; he merely published the name of a new genus there,

without listing any species. Therefore, the correct basionym was not cited
and according to ICBN Art. 33.2, the combination is invalid. The new
combination, citing the correct basionym is made herewith:

Clusia comans (Mart.) Pipoly, comb. nov. Schweiggeria comans Mart., Nov. Gen.
Sp. Pl. 3:166. 1832. Quapoya comans (Mart.) Planch. & Triana, Ann. Sci. Nat. Bot.
ser. 4, 14:239. 1860. Renngeria comans (Mart.) Meisn. ex Engl. in Mart., Fl. Bras. 12
(1):441. 1888.

Even though the generic name Schweiggeria Mart. (non Spreng.) is a later
homonym, the binomial is legitimate, as per ICBN Art. 55.1.

In the treatment of the genus Clusia for Flora of the Venezuelan Guayana,
(Pipoly et al. 1998), and subsequently (Pipoly 1997), we transferred a number
of Andean species from Oedematopus to Clusia, but did not transfer the
Andean species from Colombia and Venezuela, pending review of avail-
able material. Now that I have seen types and other material of these spe-
cies, here I transfer the remainder of the species formerly in Oedematopus,
to Clusia sect. Oedematopus, herewith.

Clusia aristeguietae (Maguire) saae comb. nov. Oedematopus aristeguietae Maguire,
Bol. Soc. Venez. Ci. Nat. 25:228.

Clusia divaricata (Cuatrec.) So comb nov. Oedematopus divaricatus Cuatrec.,
Anales Inst. Biol. Univ. Nac. Mexico 20:108. 1949.

Clusia ae ueane eames ) Pipoly, comb. nov. Ocdematopus epiphyticus Cuatrec.,
sta Acad. Colomb. Ci. Exact 8 (29):61. 1950.

ons mirandensis (Cuatrec.) aes comb. nov. Oedematopus mirandensis Maguire,
ol. Soc. Venez. Ci. Nat. 25:230.

ACKNOWLEDGMENTS

This work results from studies supported by a generous grant from the

PIPOLY AND CoGoLLo P., New species and nomenclatural notes in Clusia 409

National Geographic Society, 5575-95, for exploration of the Parque Nacional
Natural Las Orquideas, followed by the same at Reserva Natural Rio Namby.
We thank Bernardo Ramirez (PSO) for his technical assistance, as well as
the administration of FELCA (Fundacién Ecoldgica de los Colibris de Altaquer),
Marcial Bisbicuz, and Maruricio Flores. Ramiro Fonnegra, Director of HUA,
graciously made the services of Gloria Mora and Consuelo Garcia avail-
able to us to prepare the line illustrations, for which we are most grateful.
Logistical help from Juan Guillermo Ramfrez and Adriana Gémez (JAUM)
ereatly facilitated our work.

We are also very grateful to K. Gandhi (GH) for bringing the biblio-
graphic errors in the Flora of the Venezuelan Guayana treatment to the at-
tention of Pipoly. Gandhi's thorough and meticulous work is most appre-
ciated. The additional comments by D. Nicolson (US) did much to improve
the quality of the paper, for which we are most thankful.

REFERENCES

Cuoisy, J. D. 1823. Mémoi de Guttiféres et sur l’'arrangement méthodique
de cette famille. Mém. Soc. Hist. Nat. Pcs 1:210-232.
Cuarrecasas, J. 1949. Gutiferas nuevas 0 poco conocidas de Colombia. Anal. Inst. Biol.
Méx,. 22:91-112.
. 1950. Notas ala Flora de Colombia. X. Revista Acad. Colomb. Ci. Exact. 8(29):33-

64.
cae A. 1888. Guttiferae. In: C. Martius, ed. Flora Brasiliensis 12(1):382—474.
895. Guttiferae.. In: A. Engler and K. Prantl, eds. Die natiirlichen Pflanzenfamilien.
a): He 242. Verlag von Wilhelm Engelmann. Leipzig.
Ewan, J. 1951. Taxonomic notes on various species of Musaceae, Marcgraviaceae, Guttiferae
and Solanaceae of Colombia. Nat. Hist. Misc. 88:1—9.
Macuire, B. 1951. Guttiferae. In: R. Schultes, Plantae austroamericanae, VII. Bot. Mus.
Leafl. 15:55-69.
. 1977. A revision of Clusia section Cochlanthera (Choisy) Engler. Caldasia 11(55):129-

146.
Pipoty, J. 1997. Nomenclatural notes on neotropical Clusieae (Clusiaceae). Sida 17:763—
FGo:

Pipoty, J. and A. Grarr. 1995a. A synopsis of the genus Clusia sections Criuvopsis and
gs ree ae) in northern South America. Sida 16 28.
an 995b. The genus Clusia section Criuva eo in Guayana. Sida
16:649-678.
, D. Kearns, and P. Berry. 1998. Clusia. Pp. 260-294. In: P. Berry, B. Holst, and
Te eekewch. eds. Flora of the Venezuelan Guayana. Vol. 4. Caesalpiniaceae-Ericaceae.
Missouri Botanical Garden. St. Louis.
PLANCHON, J. and J. TRIANA. 1860a. Mémoire sur la famille des Guttiféres. Ann. Sci. Nat.,
Bot ser. 4, 13:307—367.
j . 1860b. Mémoire sur la famille des Guttiféres. Ann. Sci. Nat., Bot ser.
4, 14:226-367.
VESQUE, J. 1892. Ep! is Si iae ad i | tom i li
Vincennes. Paris.
= ee . Guttiferae. In: Alphonse and Casmir de Candolle, eds. Monographie
Phanetogamarum 8:1—-669. Paris.

410 Sipa 18(2)
BOOK REVIEW

JONH Lairb Farrar. 1995. Trees of the Northern United States and Canada.
1995. (ISBN 0-8138-2740-X, hbk.) Iowa State University Press, 2121
South State Ave., Ames, lowa 50014. $39.95, 502 pp, maps, line drawings,
color.

This book’s format is extremely similar to the classic Jextbook of Dendrology, by W. Harlow

& E. Harrar, pub ished in 1958. Although I found no direct reference to that work men-
tioned in the text, the 7th edition of Harlow and Harrar, published in 1991, is cited in
the Bibliography section. The author states that this book evolved from the Native Trees of
Canada, first published in 1917. Clearly, there are only so many ways to present multiple
dendrological characteristics graphically, clearly, yet with high technical accuracy, and this,
like the other aforementioned books, does just that. For each “native” species, the “pre-
ferred” English common name, latin name, French common name, its distribution, and
brief descriptions of the: leaves, buds, twigs, “seed cones” (in the case of gymnosperms),

or flowers and fruit (for angiosperms), s eetlle, vegetative reproduction, bark, wood, size
and form of bole and canopy, habitat and notes. The distrubution map, habit profile, twig
profile, leaf profile, and fruit drawing are on a page facing color photos of the cones or
flowers, ae seeds, young bark, and old bark, for most species. The entire work is di-
vided into 12 groups, the first six of which are gymnosperms (need e and scale characters)
and the second of which are angiosperms, separated by leaf morphology. While | believe
the section entitled, “Tree names” contains several important conceptual errors, they do
not con prise any vores pitfall. | would have also liked an explanation for determi-
nation of “native” species. In summary, it is one of those guides that every professional
conservationist, tie a botanist should have, along with anyone who enjoys hiking,
camping, nature walks, and other outdoor activities in the geographic area covered. The
price is more than reasonable for such an absolutely practical work, that is easy to use,
richly illustrated on acid-free paper, « and with an extra-heavy-duty binding. | am not sure
how the book would withstand a rainstorm or two, but 1 am anxious to find out, as should
everyone.—John Pipoly III

SIDA 18(2): 410. 1998

DOS NUEVAS ADICIONES A LA
ORQUIDEOFLORA MEXICANA

ADOLFO ESPEJO SERNA Y ANA ROSA LOPEZ-FERRARI

Herbario Metropolitano
Departamento de Biologia, C.B.S.
Universidad Auténoma Metropolitana-lztapalapa
Apdo. Postal 55-535
09340 MEXICO, D. E

aes@ UuUatW.INX

JAVIER GARCIA CRUZ, ROLANDO JIMENEZ MACHORRO y
LUIS SANCHEZ SALDANA
Herbario AMO
Apdo. Postal 53-123
11520 MEXICO, D.E

eric@int t COM.WX

RESUMEN
Se describen e ilustran dos ies del género Malaxis (Malaxideae, Orchidaceae)
para México, Malaxis ee y Malaxis Biaren es primera crece en el estado de Morelos
y la segunda se conoce de los estados de Morelos y Guanajuato.

ABSTRACT

Mataxis palustris and Mataxis alvaroi (Malaxideae, Orchidaceae), two new species from
Mexico, are described and illustrated. The former grows in Morelos and the latter is known
from Morelos and Guanajuato.

INTRODUCCION

El género Malaxis fue propuesto por Swartz en 1800 y comprende un
grupo de orquideas terrestres, bulbosas, ocasionalmente epifitas, que se
caracterizan por presentar una o dos hojas (ocasionalmente tres) envainadoras
y flores pequefias, verdes a verde-amarillentas, raramente purpureas dispuestas
en inflorescencias racemosas a corimbosas. El género cuenta con cerca de
300 especies de distribucién cosmopolita. Williams (1951) cita 24 especies
para México, Soto Arenas (1988) reporta 32 taxa, en tanto que Espejo &
Lépez-Ferrari (1998) registran 47. Aunque los listados mencionados nos
dan una idea del numero de representantes mexicanos del género, aun esta
lejos de completarse la revisi6n del mismo para el pais. En los ultimos 15

afios han sido descritos mas de 10 taxa (McVaugh 1985; Catling 1990; Salazar
Stipa 18(2): 411-418. 1998

412 Stipa 18(2)

1990, 1993; Salazar y Soto Arenas 1990; Todzia 1993, 1995; Gonzalez Tamayo
1991, 1992a, 1992b, 1994, 1995; Greenwood 1992) y la gran riqueza especifica
del género sugiere que aun restan por descubrirse varios mas.

Por otra parte, la distribucién de las especies de Malaxis en México es
poco conocida o en algunos casos mal entendida, debido a que la delimitacién
especifica se basa en buena medida en los caracteres florales, los cuales son
dificiles de observar 0 interpretar en el material herborizado. Ademas, la
falta de claves recientes para la identificacién de las especies conduce a que
los ejemplares, o bien no se identifiquen, o bien se determinen erréneamente.

En recientes salidas al campo a los estados de Morelos y Guanajuato, se
recolecté material de dos especies del género Malaxis, que no pudieron asignarse
a ninguno de los taxa conocidos. La revisidn bibliografica pertinente, asi
como el estudio de las colecciones del género, incluyendo material tipo,
depositadas en los herbarios AMES, AMO, ENCB, GH, MEXU, MICH,
NY, UAMIZ, VT y W nos aon ala conclusién de que se trata de dos

novedades para la ciencia, que aqui proponemos.

Malaxis palustris Espejo & Lépez-Ferrari, sp. nov. (Fig. 1). Tipo: MEXICO.
Morelos, municipio de Huitzilac, 1-2 km al W de la Laguna Zempoala, sobre el ar-
royo Las Trancas, ea ee Nacional Lagunas de Zempoala, 19° 02' 37" N, 99° 19!
20" W, 2800 m, 12 jul 1997, Espejo, 5714, Lépez-Ferrari, Garcia- Gu y Jiménez M.
(HOLOTIPO: ae ISOTIPOS: AMO, AMES).

Herba paludicola. Rizoma conspicuus. Pseudobulbus ei ovideus, inconspicuus.

Folia unica, anguste elliptica vel elliptica. Inflorescentia racemosa, pedunculo penta- alato,
racimo denso. Flores 10-30 non resupinatac. Labelum depresso-ovatum, concavum, acuminatum,
basi truncatum.

Hierba paludicola de 12-33 cm de alto. Raices pocas, originadas en la
base del seudobulbo, de 0.2—-0.5 mm de didmetro, pilosas, blanquecinas
Rizoma conspicuo, de 0.5—2.5 cm de largo por 2—3 mm de didmetro, cubierto
por las vainas. Seudobulbo hipégeo, ovoide a oblongo, inconspicuo, blanco,
de 5-10 mm de largo por 5—7 mm de ancho. Vainas dos, tubulares, estrechas,
verdosas hacia el pice, obtusas a agudas, de 3—4.5 cm de largo. Hoja unica
con una vaina tubular de 3—9.5 cm de largo que envuelve la porcién infe-
rior del escapo; l4mina de 3.5—5.5 cm de largo por 1.3—2 cm de ancho,
angostamente eliptica a elfptica, obtusa a redondeada, la base decurrente,
el margen entero, con una quilla prominente en el envés. /nflorescencia erecta,
originada de la parte apical del seudobulbo en desarrollo, racemosa, mas
larga que la hoja, con 10-30 flores sucesivas; pedtinculo de 8-16 cm de
largo por 1-1.5 mm de didmetro, alado, pentagonal en corte transversal,
racimo denso, de 2—4.5 cm de largo por ca. 1 cm de didmetro. Briacteas
florales ascendentes, verdes, triangulares, agudas, de 1—1.5 mm de largo por
ca. | mm de ancho. Flores ascendentes, de 5—6 mm de largo, amarillo-verdosas,
no resupinadas. Ovario subcilfndrico, de 2—2.5 mm de largo por ca. 1 mm

Espejo, et al., Nuevas orquideas Mexicanas 413

Gs
igs
“Vs us

2mm

Fic. 1. Malaxis palustris Espejo & Lépez-Ferrari. A. Habito de la planta; B. Detalle de la
inflorescencia; C. Flor disecada; D. Flor tres cuartos; E. Flor vista lateral; E Flor vista fron-
tal; G. Columna vista dorsal; H. Columna vista lateral; I. Columna vista ventral; J. polinios.

414 Sipa 18(2)

de didmetro. Sépalo dorsal extendido, con los margenes deflexos a revolutos,
angostamente triangular, obtuso, trinervado, de 2.5-3 mm de largo por
1.8 mm de ancho. Sépalos laterales extendidos, con los margenes deflexos a
revolutos, ang triangulares, ligeramente oblicuos, obtusos, trinervados,
de 2.5-3 mm de largo por 1.8 mm de ancho. Pétalos extendidos a recurvados,
oblongo-lineares, uninervados, de 2.5—3 mm de largo por ca. 0.3 mm de
ancho, el dpice redondeado. Labelo depreso-ovado, abruptamente acuminado,
céncavo, abrazando la columna en posicién natural, con una quilla media
longitudinal evidente de color verde oscuro, de 2.5 mm de largo por 2.2
mm de ancho, la base truncada. Co/umna corta, cuadrada, comprimida
dorsiventralmente, truncada, de 0.7 mm de largo por 0.7 mm de ancho.
Antera transversalmente oblonga, bilocular, de 0.2 mm de largo por 0.6
mm de ancho. Polinarios dos, separades, divergentes, cada polinario formado
or dos polinios fusionados entre sf; cada par de polinios obpiriforme, oblicuo,
comprimido dorsiventralmente, amarillo, de 0.3 mm de largo con un viscidio
apical, formado por una gota viscosa, de color Ambar. Est/gma ventral, semicircular,
concavo. Capsula elipsoide, de 4-5 mm de largo por 2 mm de didmetro.

Material examinado: México. Morelos: municipio de Huitzilac, 1-2 km al W de la Laguna
peer sobre el arroyo Las Trancas, Parque Nacional Lagunas de Zempoala, 19° 02’
37" N, 99° 19’ 20" W, 2800 m, 12 jul 1997, Garcta-Cruz 751, Jiménez M., Sanchez S. y
Tae O. (AMO, UAMIZ).

Etimologta.—el nombre de la especie hace alusion a su habito paludicola
ya que crece en Ilanos permanentemente inundados rodeados por bosques
de Abies.

Malaxis palustris se conoce hasta el momento sélo de la localidad tipo,
en el estado de Morelos, sin embargo, es de esperarse su presencia en el
vecino municipio de Ocuilan, en el Estado de México. La preferencia de
las plantas de esta especie por los lugares abiertos y anegados, habito poco
comun en el género, la distinguen del resto de las especies mexicanas. Otros
caracteres peculiares de M/. palustris son el rizoma evidente, las flores no
resupinadas y el labelo depreso-ovado, abruptamente acuminado en el apice
y truncado en la base. El conjunto de todas estas caracteristicas distinguen
inconfundiblemente aM. palustris de cualquier otra especie mexicana conocida.

Cabe sefalar que en los alrededores de la localidad tipo de M. palustris, crecen

también M. ehrenbergti (Rchb. f.) Kuntze, M@. myurus (Lindl.) Kuntze, ©.
salazarit Catling, M. souler L. O. Williams, M. streptopetala (B.L. Rob. &

reenm.) Ames y M. tenuis (S. Watson) Ames, sin embargo ninguna de
ellas se encuentra en lugares pantanosos, sino que preneiee sitios elevados
y secos dentro de los bosques de pino y/o Abies.

Malaxis alvaroi Garcia-Cruz, R. Jiménez & L. Sanchez, ] nov. (Fig. 2).
TIPO: MEXICO. MORELOS: municipio de Tepoztlan, sobre la via del ferrocarril México

Espejo, et al., Nuevas orquideas Mexicanas 415

Cuernavaca, al § de San Juan Tlacotenco, 19° 01' 00" N, 99° 06' 48" W, 2250 m, 27
jul 1996, Jiménez M. 1962, Garcta-Cruz, Lépez-Ferrari y Espejo (HOLOTIPO: AMO;
ISOTIPOS: ENCB, IEB, UAMIZ)

Herba terrestre. Rizoma ausente. Pseudobulbus hipogeus, ovideus. Folia unica, ovata,
apice obtusa vel rotundata, basi leviter cordata. Inflorescentia el corimbosa, pedunculo
deca-alato, racimo denso. Alas minutae erosae vel serratae. Flores 20-45, resupinatae. Labelum
deltato- sagitatum, leviter concavum, acutum, basi bilobulatum. Lobuli triangularis nonunquam

apice falcati.

Hierba terrestre de 12—23 cm de alto. Raéces pocas, originadas de la base
del seudobulbo, de 0.3—0.6 mm de didmetro, pilosas, blanquecinas. Rizoma
ausente. Seudobulbo hipdgeo, ovoide, suculento, blanco, de 7 mm de largo

or 4.3—8 mm de ancho. Vaina una, tubular, estrecha, verdosa, obtusa a
redondeada, de 2—4 cm de largo. Hoja unica con una vaina tubular, de 2.5—
3.5 cm de largo, ligeramente comprimida, con cuatro quillas evidentes que
envuelve la porcidén inferior del escapo; lamina de 3—4.5 cm de largo pe
1.8—2.5 cm de ancho, ovada, obtusa a redondeada, la base li :
el margen levemente ondulado, sobre todo hacia la Gace! con cinco venas
ligeramente hundidas en el haz y prominentes en el envés, la vena media
marcadamente quillada, de color verde oscuro. /nflorescencia erecta, originada
de la parte apical del seudobulbo, racemosa a corimbosa, mas larga que la
hoja, con 20—45 flores sucesivas; pedtinculo de 4-10 cm de largo por 0.8—
1.5 mm de didmetro, con diez alas diminutamente aserradas a erosas, racimo
denso, de 1.5—2.5 cm de largo por 0.8-1.5 cm de didmetro; raquis alado,
las alas diminutamente aserradas a erosas. Bracteas florales extendidas, verdes,
triangulares, agudas, de 0.5 mm de largo por 0.5 mm de ancho. Mores ascendentes,

—4.2 mm de largo, verde oscuras, resupinadas. Ovario torcido, quillado,
subcilindrico, de 1.7—4 mm de largo por 0.3—0.5 mm de didmetro. Sépalo
dorsal extendido, ee ovadoa neces: obtuso, trinervado, de 1.5-—
2,2 mm de largo por 1—1.3 mm de ancho , brevemente
connatos en la base, aplanados, aes oblicuos, ovados a lanceolados,
obtusos, trinervados, de 1.6—2.2 mm de largo por 1-1.3 mm de ancho.
Pétalos recurvados, lineares, uninervados, de 1.5—2 mm de largo por ca.
0.2 mm de ancho, el pice redondeado. Labelo deltado sagitado, ligeramente
concavo, pentanervado, de 1.7—2.5 mm de largo por 2.1—2.3 mm de ancho,
agudo, la base con dos Idbulos laterales triangulares con el dpice en ocasiones
falcado. Columna corta, subcuadrada, comprimida dorsiventralmente, truncada,
de ca. 0.5 mm de largo por ca. 0.5 mm de ancho. Antera dorsal, bilocular,
reniforme, de 0.2 mm de largo por 0.4 mm de ancho. Polinarios dos, separados,
divergentes, cada polinario formado por dos polinios fusionados entre si,
obovoide, oblicuo, amarillo

cada par de p
de 0.3 mm de largo con un viscidio Aertel formado por una gota viscosa de
color dmbar. Estigma ventral, subcuadrado, céncavo. Capsula obovoide,
ascendente, de 5 mm de largo por 2 mm de didmetro.

416 Sipa 18(2)

5 ay
aR

Imm
G

(are 4
lmm

Fic. 2. Malaxis alvaroi Garcia-Cruz, R. Jiménez & L. Sanchez. A. Habito de la planta; B.
Detalle de la inflorescencia; C. Flor disecada; D. Flor vista lateral; E. Flor vista frontal; E
Columna vista dorsal; G. Columna vista lateral; H. Colummna vista ventral; I. Polinios.

Espejo, et al., Nuevas orquideas Mexicanas 417

Material examinado: MEXICO. Ee municipio de San Luis de la Paz, camino
de Las Mesas de Jestis al Vergel, 21° 31' 59" N, 100° 25' 56" W, 2240 m, 9 ago 1996,
Sanchez S. 375, Jiménez M. y Garcia-Cruz ave UAMIZ). Morelos: municipio de Tepoztlan,
km 93-94 sobre la via del ferrocarril México-Cuernavaca, al S de San Juan Tlacotenco, 19°
01' 00" N, 99° 06' 48" W, 2250 m, 27 jul 1996, Espejo 5430 y ee (AMO, UAMI/Z).

Etimologia.—el epiteto asignado a este taxon se dedica al bidlogo Alvaro
Flores Castorena, amigo y colega con quien hemos recorrido gran parte del
estado de Morelos recolectando orquideas y otras plantas.

Esta especie crece en los bosques de encino secos y generalmente forma
colonias pequenas dificiles de encontrar.

Malaxis alvaroi se reconoce por sus flores pequefias de menos de 4.3 mm
de largo y de color verde obscuro, dispuestas apretadamente en una inflorescencia
racemosa a corimbosa, que se va alargando por la permanencia de las flores
marchitas en la parte inferior de la misma. Otros caracteres distintivos son
el labelo deltado sagitado, con los lébulos basales agudos y las alas aserradas
a erosas del pedunculo y el raquis.

Por sus caracteres vegetativos y florales, como son la hoja unica y cordada,
la inflorescencia racemoso-corimbosa y el labelo triangular y sin ornamentaciones,
Malaxis alvaroi puede ubicarse dentro del grupo de especies formado por
M. brachystachys y M. corymbosa. En el cuadro 1 pueden apreciarse las
caracteristicas comparativas entre éstas (Tabla 1).

Tasia 1. Comy cién de alg isti M. alvaroi, M. brachsytachys, y M. corymbosa

Caracteristicas M. alvarot M. brachsytachys M. corymbosa

| itud d {ui | 4-10 3-7 6.5-9.5

orn neha as peduinculo aserradas a erosas alas lisas alas lisas
inflorescencia corimbo alargado imt to y aplanad orimbo co aplanadc
| 1 de las flores ( ) 3-4.2 9.5-12 6-8.5

longitud a res (mm) ae ee

flor

labelo

Varn

color de las flores verde obscuro, opacas amarillo verdoso, hialinas amarillo verdoso, hialinas
] Fe ae pa ee 1.7-4 10-12 6-7

418 Stipa 18(2)
AGRADECIMIENTOS

Este trabajo fue patrocinado parcialmente por la Comision Nacional ae
el Conocimiento y Uso de la Biodiversidad PON eB.) a trayes de los
proyectos G016, H043 y H098. Deseamos hacer f imiento
a Victoria Sosa, Kerry A. Barringer, Lawrence K. Magrath, Eric Hagsater y
Gerardo Salazar por la revisibn critica del manuscrito. Asimismo agradecemos
a los curadores de los herbarios consultados por las facilidades brindadas
para la revisién del material. Los dibujos fueron elaborados por Rolando
Jiménez Machorro.

REFERENCIAS

CATLING, PM. 1990. Malaxis salazarti, a new species from Mexico and Northern Mesoamerica.
Orquidea (Mexico City) 12:93-10

Espryo, A. and . LOPEZ-FERRARI. (1998). Las Monocotiledéneas Mexicanas una Sinopsis
Floristica 1 iG ista de Referencia PARTE VII. Orchidaceae 2. Consejo Nacional de la
Flora Mee , A.C., Universidad Auténoma Metropolitana-Iztapalapa, México, D.E.

GONZALEZ TAMAYO, R. 1991. Malaxis martinezii (Orchidaceae), una especie nueva muy
cercana a M. reichei. Bol. Inst. Bot. Univ. Gauadalajara 1:33-37.

GONZALEZ Tamayo, R. 1992. Malaxis sree (Orchidaceae), una especie nueva en honor
de una estudiosa de la Botanica. Bol. Inst. Bot. Univ. Gauadalajara 1:88—‘

GONZALEZ Tamayo, R. 1992. Malaxis ruizii, una nueva especie del oceidente de México
(Orchidaceae). Bol. Inst. Bot. Univ. Gauadalajara 1:211-216.

GONZALEZ Tamayo, R. 1994. Una curiosa Malaxis del occidente de México. Acta Bot. Mex.

9 Q: am ¢ 9

—

(GONZALEZ Ta AMAYO, R. 1995. Malaxis jolla Sepals ae), interesante especie del occidente
de México. Bol. Inst. Bot. Univ. Guadalajara 128.
REENWOOD, EW. 1992. Malaxis She a new species from Oaxaca. Orquidea (Mexico
City) 12:199-203.
McVaucu, R. 1985. Flora Novo-Galiciana. Vol. 16. Orchidaceae. University of Michigan

Press. Ann Arbor

G

SALAZAR, G.A. 1990. Malaxis hagsateri, una nueva especie de Guerrero, México. Orquidea
(Mexico vaste 2:81-

SALAZAR, G.A. 1993. Malaxis wendtii, a new oe species from Coahuila and Nuevo Leén,
Mexico. Slat (Mexico City) 13:281-

SALAZAR, G.A. and M.A. Soro Arenas. 1990. Una nueva especie de Malaxis (Orchidaceae)

e flores oe norte de Chiapas. Acta Bot. Mex. 10:45—49,

ne Arenas, M.A, | 8. Listado actualizado de las orquideas de México. Orquidea (Mexico
City) 11:23 coed

Topzia, C. 1993. Malaxis hintonii, a new species from Northern Mexico. Orquidea (Mexico
City) 13:121-124.

TODA; C1995: ra chica a new species of Orchidaceae from Nuevo Leén, México,
Brittonia 47:2

WittAMS, L.O. 1951. {hc Orchidaceae of Mexico. Ceiba 2:1—321

SEEDLING DEVELOPMENT IN SPECIES OF
CHAMAESVCE PUPHORBIACEAR WATR-LERECT
GROWTH HABITS

W. JOHN HAYDEN and OLGA TROYANSKAYA

Department of Biology
University of Richmond
Richmond, VA 23173, U.S.A.
ABSTRACT
Seedling development is described for Chamaesyce hirta, C. hypericifolia, and C.
mesembrianthemifolia as discerned by light seg? and ae electron microscopy.
l xh these species ultimately develop erect to ascending growth habits, epicotyl de-
velopment is limited to the production of a ok pair of leaves oe immediately superjacent
to and decussate with the cotyledons. The shoot system develops from one or more buds
located in the axils of the cotyledons. In all respects, seedling ontogeny is very similar to
that of previously studied prostrate species of Chamaesyce. Evidence from seedling ontog-
eny thus contradicts a hypothesis concerning homologies of plant form pertinent to the
origin of Chamaesyce from Euphorbia that was first sitieuilated by Roeper in 1824. These
results support an alternative hypothesis based on proliferation of branches from the coty-
ledonary node in hypothetical ancestral elements within Euphorbia where this morphol-
ogy can be found in perennial hemicryptophytes as well as certain annual species.

—

RESUMEN

e describe el desarrollo de la semilla de ae iene C. eee gon
Be iri oH al microscopio éptico y micr le barrido. Aunque
estas especies desarrollan finalmente habitos de pee ie te erectos a Re ae el
desarrollo del epicétilo se limita inicamente a la produccién de un par de hojas localizadas
inmediatamente encima y decusadas con los cotiledones. El sistema de ramas se desarrolla
a partir de una o mds yemas localizadas en las axilas de los cotiledones. La ontogenia de la
semilla es en todos los aspectos muy semejante a la de las especies de Chamaesyce postradas
estudiadas previamente. Esta evidencia de la ontogenia de la semilla contradice la hipotesis
peas a us ance de la forma de la planta, relativas al origen de Chamaesyce a partir
de £ or Roeper en 1824. Estos resultados apoyan la hipotesis alternativa
basa en la prliferacin ee eae paa del nudo de los cotiledones en los hipotéticos

morfologfa en hemicriptéfitos

perennes asf como en algunas especies anuales.

The genus Chamaesyce Gray can be distinguished from its close relatives
in Euphorbia L. by a series of morphological, physiological, and develop-
mental characters (Webster 1967, 1994; Koutnik 1984). The peculiar fea-
tures of seedling ontogeny in Chamaesyce are often asserted to be charac-

teristic for the genus (Degener & Croizat 1938; Koutnik 1987). However,

Stipa 18(2): 419-431. 1998

420 Sipa 18(2)

seedlings of only a few species have been studied anatomically (Veh 1928;
Mangaly et al. 1979; Rosengarten & Hayden 1983; Hayden 1988). More-
over, these few anatomical studies have generally focused on species of sec-
tion Chamaesyce, a group characterized by radiating, prostrate branches.
This paper describes seedling development in species with erect to ascend-
ing growth habits for comparison with the known structures and develop-
mental events of prostrate species. Further, this paper critically assesses the
interpretation of Mangaly et al. (1979) who described extra-axillary origin
of lateral branching in seedlings of Chamaesyce and it is the first study to
examine Chamaesyce seedlings with the scanning electron microscope (SEM).
We document and describe seedling development for three species rep-
resenting two sections of the genus characterized by erect to ascending stems.
Chamaesyce hyperic ifolia (L.) Millsp., the type species of section Hypericifoliae
(Boiss.) Hurus., is a widely distributed weed (Holm et al. 1979) thought
originally to be native to warm regions of the Americas (Webster 1967;
Koutnik & Huft 1990). Chamaesyce hirta (L.) Millsp. is also classified in
section Hypericifoliae and has similar present day status as a pantropical
weed (Cardenas et al. 1972; Holm et al. 1979); however, it may be native
to both the New and the Old World tropics (Koutnik & Huft 1990). Chamaesyce
mesembrianthemifolia (Jacq.) Dugand is a shrubby plant classified in sec-
tion Sclerophyllae (Boiss.) Hurusawa and found near seashores of the Car-
ibbean and northern South America (Long & Lakela 1976; Acevedo-Rodriguez
1996). Although C. Aypericifolia is often characterized as an herb, under
favorable conditions it can produce weakly woody stems approaching a meter
in height and | cm in basal diameter; thus, it can attain a stature compa-
rable to that of C. mesembrianthemifolia. In contrast, C. hirta is smaller,
and somewhat intermediate between the prostrate growth habit typical of
section Chamaesyce and the erect growth habits of sections Hypericifoliae
and Sclerophyllae. In C. hirta, each plant produces multiple stems, but sel-
om as many as found in species of section Chamaesyce, and while these
stems are sometimes more or less prostrate, they more frequently ascend,
but seldom to heights exceeding 15 or 20 cm.

MATERIALS AND METHODS

Specimens of C. Airta were collected in 1982 from weeds among nurs-
ery stock in a Richmond, VA, garden center. Plants of C. mesembrianthemifolia
were collected at West Summerland Key, Florida, in 1983 and plants and
seeds of C. hypericifolia were collected from Big Pine Key, Florida, in 1986.
All three species have been maintained subsequently in greenhouse culti-
vation at the University of Richmond. Adult plants were prolific in culti-
vation; over several years C. hypericifolia produced many thousands of seedlings
whereas C. /irta and C. mesembrianthemifolia each produced several hun-

HAYDEN AND TROYANSKAYA, Seedling development in Chamaesyce 421

dred. Preparations of C. hirta and C. hypericifolia were derived from seed-
lings obtained spontaneously from containers of sterile soil placed in the
vicinity of mature greenhouse-grown plants. Preparations of C.
mesembrianthemifolia were derived largely from a dense mass of seedlings
collected in the wild from the surface of a small anthill and supplemented
with greenhouse materials. Herbarium vouchers at URV for materials studied
include: C. hirta—Hayden 614; C. mesembrianthemifolia—Hayden 709, and
C. hypericifolia—Hayden 1433, 3252 and 3281. All specimens were pre-
served in FAA (formalin—acetic acid—70% alcohol). Numbers of seed-
lings studied for each species are as follows: C. irta—30 for light micros-
copy (LM); C. Aypericifolia—30 for LM and 42 for SEM;
mesembrianthemifolia—\7 fe LM and 14 for SEM.

For light microscopy, fixed seedlings were trimmed to remove structures
more than 5 mm below or above the cotyledonary node (if present), dehy-
drated in a tert-butanol series, embedded in paraffin, sectioned at 10 pm,
affixed to slides with Bissing’s adhesive (Bissing 1974), and stained in a
combination of safranin and hematoxylin (Johansen 1940). Photomicro-
graphs were prepared from Kodak Technical Pan film developed in Kodak
HC110 developer at dilution F. For SEM, fixed seedlings were trimmed,
dehydrated in ethanol, subjected to critical point-drying with CO as the
intermediate solvent, affixed to stubs with aluminum tape, sputter-coated
with a gold-palladium mixture to a thickness of 40 nm, and observed with
a Hitachi S-2300 SEM. Scanning electron micrographs were prepared from
Kodak Tri-X film developed in Kodak HC110 developer at dilution B.

RESULTS
External morphology

In the seed and during initial stages of germination, the cotyledons are tightly
appressed. Upon germination, blades of the cotyledons diverge first, leav-
ing their petioles in close contact. Soon, however, petioles also diverge, re-
vealing primordia for the first pair of true leaves (Figs. 1-3). Following Hayden
(1988), these first leaves are referred to as v-leaves to indicate their sup-
posed homology with leaves on the vegetative axis of Euphorbia species classified
in subgenus Agaloma (Raf.) House and subgenus Esu/a Pers. Upon their
full expansion, v-leaves are inserted directly superjacent to and decussate
with the cotyledons (Fig. 3). There is no residue of meristematically active
cells at the epicotyl apex following v-leaf formation (Figs. 4, 9, 10, 15-17).
Seedling growth continues via lateral branches that arise from buds axil-
lary to the cotyledons (Figs. 4-8, 10-12, 15, 17, 20).

Although multiple buds routinely develop in the axils of each cotyledon
(Figs. 11, 20), just a single lateral branch dominates early growth in C. hypericifolia
and C. mesembrianthemifolia (Figs. 5, 6). This first, dominant branch is

422 SIDA 18(2)

Fics. 1—4 eralings of C Relies e, SEM. C. hypericifolia, Hayden 328 1, v-leaf ae
dia, top view of seedling soon after divergence of cotyledonary petioles; bar = 100 pm. 2.
oe hyper ictfolta, Hayden. 3690, seledone and v-leaf primordia, lateral view; bar = 250 um.
9 4G. (ie Aa loa Hayden 709. 3. Cotyledons and v-leaf primordia, lateral
view; bar = 500 pm picotyl apex, petioles of cotyledons and y-leaves, and cotyledon-

ary axillary buds; Gaee 100 um, c = cotyledon; v = v-leaf.

erect or slightly inclined from vertical. Growth of additional lateral branches
at the cotyledonary node as plants grow older is highly variable in these
two species. Vigorous specimens with sparse or no competing nearby veg-
etation tend to produce one or two additional basal branches, but these
remain smaller than the first branch unless the latter is removed or dam-

HayDEN AND TROYANSKAYA, Seedling development in Chamaesyce 423

Fics. 5,6. Cae ion node of aie of one meembvonbenfli foe
709, each with one v-leaf removed, SEM. 5. Bud in axil of cotyledon; b n. 6.
Young lateral branch, arising from axil of cotyledon; bar = 500 pm. c = seca V=Vv-
leaf.

aged. It is not unusual for nearly the entire aerial system in these species to
develop from the first dominant branch that arises at the cotyledonary node.

Initially, seedlings of C. Airta are similar to those of C. hypericifolia and
C. mesembrianthemifolia. However, in C. hirta, multiple lateral branches
develop from the cotyledonary node. The first two branches arise one each
from the axils of the cotyledons and, frequently, two additional branches
develop from buds located at the bases of the first two branches.

Anatomical structure

Cotyledon stage.—The hypocoty] is traversed by four vascular bundles that
ascend from the radicle to the cotyledonary node. Two of these bundles,
each located on opposite sides of the axis, constitute the median traces to
the cotyledons; each passes directly from the hypocotyl into the petiole of
its respective cotyledon. The other two bundles form four lateral traces to
the cotyledons; each splits into two bundles at the cotyledonary node and
the resultant pair of traces diverge towards cotyledons on opposite sides of
the stem. Cotyledonary node vasculature is thus trilacunar with split later-
als (Fig. 19). The system of non-articulated branched laticifers that ulti-
mately permeates the plant body arises from initials located external to the
vascular tissue at the cotyledonary node.

V-leaves.—V-leaves arise on the flanks of the epicotyl apex (Figs. 1, 9).
As soon as v-leaf primordia can be detected, cells of the epicotyl apex are
larger and more vacuolate than those of the v-leaf primordia (Fig. 9). Thus,
the only meristematically active cells of the epicotyl apex are fully consumed
in formation of the v-leaves. Each v-leaf is vascularized by three traces that
differentiate from procambium near the split lateral traces to the cotyle-
dons; of the three traces for a given v-leaf, two traces insert on one side of
the cotyledonary split lateral and one trace inserts on the opposite side. General

24 Sipa 18(2)

Fics. 7, 8. Seedlings of Chamaesyce hirta, Hayden 614, LM. 7. Cotyledons, their lateral
buds, and portions of one v-leaf, longitudinal section in the cotyledonary plane; bar = 50
pm. 8. Cotyledon petioles, their lateral buds, and v-leaf petioles; bar = 100 pm. c = coty-

ledon; v = v-leaf.

vascular development of the seedling continues as the v-leaves develop. Once
the v-leaves are fully expanded, the hypocotyl contains four well-defined
vascular strands that supply the first four leaves of the seedling.

Branch development.—SEM micrographs of intact seedling apices and
LM sections reveal branch primordia in the axils of the cotyledons (Figs. 4,
7,8, 12, 15, 17, 20). Lateral branches arise from ordinary lateral buds lo-
cated at the cotyledonary node. The densely-stained and meristematically
active lateral buds (or the subsequent active branch apex, e.g., Fig. 13) contrast
sharply with the inert epicotyl apex (Figs. 7, 8, 10, 12, 17). Each axillary
bud is also associated with a non-vascularized, persistent stipule-like flap
of tissue (Figs. 10, 11, 18). Whereas stipules located on subsequent nodes
(1.e., nodes of the lateral branch) generally consist of a planar interpetiolar
sheath, stipules at the cotyledonary node are elongate and bear a distinct
curve that conforms with the curved surface of its associated bud
primordium.

At their initiation, the buds that produce lateral branches are clearly not
aligned with the axis defined by the hypocotyl and truncated epicoryl. In
time however, the first, dominant, branch of fully erect species assumes an
apparent axial position (Fig. 14), a consequence of its growth and expan-
sion concomitant with that of the hypocotyl.

HAYDEN AND TROYANSKAYA, Seedling development in Chamaesyce 425
DISCUSSION

Based on previous literature (Veh 1928; Mangaly et al. 1979; Rosengarten
& Hayden 1983; Hayden 1988) and the species described above, early stages
of seedling development in Chamaesyce appear to be similar in sections
Chamaesyce, Hypericifoliae, and Sclerophyllae. Common features include the
vascular architecture of the cotyledonary node, the development of v-leaves
and their vascular supply, and the origin of lateral branches from the axils
of the cotyledons. The potential for multiple branches from the cotyledonary
node is also uniform throughout the species for which seedling ontogeny
is known. The chief differences between previously studied prostrate spe-
cies and the erect or semi-erect species documented here involves the num-
ber of lateral branches that develop from the axils of the cotyledons and
their orientation with respect to gravity. Thus, species of section Chamaesyce
routinely produce multiple branches that radiate at ground level, whereas
most species of sections Hypericifoliae and Sclerophyllae produce one dominant,
erect branch and, sometimes, another small, subsidiary branch. Chamaesyce
hirta appears somewhat intermediate in that it produces a limited number
of semi-erect branches.

Mangaly et al. (1979) reported on seedling development in Chamaesyce
hirta and C. thymifolia (L.) Millsp. We find their illustrations consistent
with the anatomy and morphology of the species reported here and in Hayden
(1988). We differ, however, in the interpretation of certain aspects of seed-
ling structure and developmental processes. For example, Mangaly et al.
(1979) failed to notice the absence of an epicotylar apical meristem upon
development of the v-leaves. Thus, they interpreted the first lateral branch,
which actually develops from the axil of a cotyledon, to be the “main axis”
and they also described the second lateral branch, which develops from the
axil of the other cotyledon, as “extra-axillary.” On the basis of gross exter-
nal morphology, numerous authors have expounded on the significance of
the absence of true epicotyl development (main axis) in Chamaesyce (e.g.,
Wheeler 1941; Degener & Croizat 1938; Webster 1967). Both the absence
of epicotyl and the axillary origin of the first two branches are clearly indi-
cated by the LM and SEM evidence presented here. Hayden (1988) earlier
refuted other evidently erroneous ontogenetic interpretations of seedling
structure in Chamaesyce found in Veh (1928) and Degener and Croizat (1938).

We hypothesize that seedling form in Chamaesyce is derived from plants
with growth habits that are widespread in Euphorbia subgenus Agaloma and
subgenus Eswla. Such plants produce ordinary, vegetative stems from the
seedling epicotyl that eventually terminate in a single cyathium followed
by a pleiochasial and dichasial pattern of cyathium production; they also
produce additional vegetative axes from the cotyledonary node which also

become pleiochasial or dichasial (Fig. 21-A). Euphorbia helioscopia L., as

426 Sipa 18(2)
illustrated in Korsmo (1954; fig. 111) or Holm et al. (1997; fig. 41-2),

provides a good example of this growth habit in an annual species. Reduc-
tion of the epicotyl to the first pair of leaves and accelerated proliferation
of branches from the cotyledonary node (Fig. 21-C) would result in the
growth habit found in Chamaesyce. By this interpretation, the majority of
the shoot system in Chamaesyce would be homologous with proliferative
cotyledonary branches in species of Euphorbia.

The hypothesis of homology at the cotyledonary node described above
stands in partial contradiction to the oft-cited hypothesis of Roeper (1824)
(Fig. 21-A,-B,-D) who said that foreshortening of the epicotyl results in
development of pleiochasial branches (inflorescence axes) at ground level
(see, for example, Wheeler 1941; Webster 1967). Pleiochasial branches in
Euphorbia arise individually from the axils of a whorl of leaves at the apex
of the epicotyl-derived main axis of the plant (Hayden 1988). Iflateral branches
in seedlings of Chamaesyce developed also from the axils of the v-leaves,
then Roeper’s hypothesis could be supported. However, this paper provides
additional confirmation of the fact that lateral branches in Chamaesyce are
strictly axillary to the cotyledons and never arise from axils of the v-leaves
(Hayden 1988).

Plants with well-developed epicotyls and proliferative branches from the
cotyledonz ary node, the hypothesized condition in ancestors of Chamaesyce,
are common in Euphorbia. Vhis growth habit occurs in both hemicryptophyuc
perennials and annuals. Among the hemicryptophytic forms, some famil-
iar garden examples from subgenus sua include Euphorbia myrsinites L.
and E. epithymoidesL. (E. polychroma A. Kern.); Euphorbia corollata L., widespread
in eastern North America, and its close relatives in subgenus Agaloma sec-
tion Tithymalopsis (Klotzsch & Garcke) Boiss., also proliferate from the
cotyledonary node. Examples of annual species with proliferative branches
from the cotyledonary node include FE. helioscopia from subgenus Esula, EF.
exstipulata Engelm. from subgenus Agaloma, and E. dentata Michx. from
subgenus Poinsettia (Graham) House. In Chamaesyce, ane with prolif-
erative cotyledonary nodes include hemicryptophytic perennials (Simmons
& Hayden 1997), prostrate annuals (Hayden 1988), erect to ascending annuals
(present study), sub-shrubs (present study), and small trees (Koutnik 1987).
In contrast to the situation in Chamaesyce, Euphorbia species possess well-
developed epicotyls.

Despite the uniformity of form and development that seems to be emerging
from studies of Chamaesyce seedlings, examination of a few additional critical
taxa appears warranted. Based on our unpublished observations of very limited
material, seedlings of C. acuta (Engelm.) Millsp. appear to possess a rela-
tively normal pattern of seedling development, with well-developed epico-
tyls. Chamaesyce acuta also lacks C4 photosynthesis (Webster et al. 1975)
and its inclusion within Chamaesyce might thus be doubted. Confirmation

HAYDEN AND TROYANSKAYA, Seedling development in Chamaesyce 427

Fics. 9-14. Seedlings of Chamaesyce hypericifolia, LM; 9, 13, Hayden 3252; 10-12, 14,
Hayden 3281. an Petioles of cotyledons and v-leaf primordia, cross section at the epicotyl
apex; bar = 50 pm. 10. Petioles of cotyledons, their lateral buds, and petioles of fully ex-
panded ee cross section at the epicotyl apex; bar = 200 pm. 11. Same seedling as Fig.
10, cross section just above insertion of cotyledons; bar = 00 1 joa 2 Beco of cotyle-
as and their axillary buds, longitudinal section in the cotyledonary plane; bar = 200 pm
of lateral branch, longitudinal section; bar = 100 pm. 14. Cotyledonary node of
sh dominated by one lateral branch, longitudinal section in the intercotyledonary
slane; bar = 200 jm. b = branch from axil of cotyledon, ¢ = cotyledon, h = hypocotyl, s =

Pp
stipule-like flap, v = v-leaf.

Stipa 18(2)

+)
7)
htt
1
4
ry

Fics. 15-20. Seedlings of Chamaesyce mesembrianthemifolia, Hayden 709, LM. 15. Petioles
of cotyledons, their lateral buds, and petioles of v-leaves, cross section below the epicoty!
pex; bar = 50 pm. 16. v-leaf primordia, longitudinal section in the intercotyledonary plane;
bar = 100 jim. 17. Petioles of cotyledons and their lateral buds, longitudinal section in the
cotyledonary plane; bar = 200 pm.

and adjacent petiole bases of cotyled

is)

c

8. Base of lateral branch arising from cotyledon axil
ons and v-leaves, cross section at the epicotyl apex; bar
200 pm. 19. Coryledonary node from seedling with expanded v-leaves, note median traces
gle arrows) and split lateral traces (double-headed arrows) to the cotyledons; bar = 100
yim. 20. Petioles of cotyledons, their lateral buc

is, and petioles of v-leaves; bar = 100 pm. b
branch from axil of cotyledon, ¢ = cotyledon, s = stipule-like flap, v = v-leaf.

HAYDEN AND TROYANSKAYA, Seedling development in Chamaesyce 429

Fic, 21. Comparison of Roeper’s (1824) and Hayden’s (1988) hypotheses concerning the
origin of the plant body in Chamaesyce. A. Hypothetical ancestor with habit common in
Euphorbia subgenus Agaloma or subgenus Esula, i.e., epicotyl well-developed, with termi-
nal cyathium: n system and reiterative lateral branches from the cotyledonary
node! ly Eypornecaal i rcrmiediate (Roeper’s Hypothesis) with reduced epicotyl develop-
ment and no reiterative lateral branches. C. Hypothetical intermediate (Hayden’s Hypothesis)
with epicotyl aborted above the first node. D. Erect and prostrate growth habits in Chamaesyce.

of seedling development in this and supposedly related species of section
Acutae (Boiss.) Webster (see discussion in Mayfield 1991) would thus prove
useful. Also, seedling development in C. potentilloides (Boiss.) Croizat [=C.
caecorum es ex Boiss.) Croizat] of southern Brazil and adjacent regions
Is letely unknown; because this is a hemicryptophyte species that routinely

produces picechasial inflorescences reminiscent of the reproductive branches

in subgenus Agaloma and Esula of Euphorbia (Simmons & Hayden 1997),
a developmental study of its seedlings may provide useful phylogenetic insight
for the genus as a whole. Finally, certain Hawaiian Chamaesyce species en-
demic to the island of Kauai appear to have ordinary seedling development
with well-developed epicotyls (Koutnik 1987); anatomical comparisons between
these epicotyl-present and epicotyl-absent species of Chamaesyce should prove

critical in assessing relationships among the Hawaiian species of Chamaesyce.
A developmental mutant known in Arabidopsis Heynh. (Brassicaceae)
results in a pattern of epicotyl deletion reminiscent of this hallmark feature

430 Sipa 18(2)

of seedling ontogeny in Chamaesyce. In Arabidopsis, plants homozygous for
the WUSCHEL (WUS) allele form a pair of leaves above the cotyledons
but fail to develop any other ordinary epicotylar structure (Clark 1997). In
WUS mutants, the epicotyl apex remains somewhat flattened and essen-
tially non-meristematic, although it can initiate adventitious meristems that
reiterate the abruptly terminated structure of these mutant seedlings (Clark
1997). Evidently, ordinary branches do not form at the cotyledonary node
in these WUS mutants of Arabidopsis, so the analogy with developmental
events in Chamaesyce is only approximate.

—

ACKNOWLEDGMENTS

This study was supported by the Undergraduate Research Committee
and Kuyk Chair funds from the University of Richmond, and by a grant
from the Jeffress Memorial Trust. Marilynn Rosengarten, Thomas Felts,
and Susie Kim assisted in the preparation of microscope slides. Sheila Hayden
assisted in the production of Fig 21.

REFERENCES
ACEVEDO-RODRIGUEZ, P. (and col oo 1996. Flora of St. John U.S. Virgin Islands.
n. New York Bot. Gard. Vol

BissiNG, D.R. 1974. Haupt’s ee ie mixed with formalin for affixing paraffin
sections to slides. Stain Technol. 49:116-117.

Cardenas, J., C.E. Reyes, and J.D. Do... nae ‘Tropical weeds, malezas tropicales. Vol.
1. Instituto Colombiano Agropécurio, Bogot:

Ciark, S.E. 1997. Organ formation at the oe shoot meristem. Pl. Cell 9:1067—
1076.

Decener, O. and L. Croizar. 1938. Chamaesyce. In: O. Degener, Flora Hawaiiensis. [Offset
printed in 1946 with some revisions, not paginate

Haypen, W.J. 1988. Ontogeny of the cotyledonary region of Chamaesyce maculata
(Euphorbiaceae). Amer. J. Bot. 75:1701-1713.

Hou, L., J. Dott, E. How, J. PANCHO, and J. HERBERGER. 1997. World weeds, natural
histories and distribution. John Wiley & Sons, New York.

Hou, L., J.V. PANCHO, J.P. HerBeRGER, and D.L. PLucKNETT. 1979. A geographical atlas
of world weeds. John Wiley & Sons, New York.

JOHANSEN, D.A. 1940. Plant ee is McGraw-Hill Book Co., New York.

Korsmo, E. 1954. Anatomy of weeds. Grondahl & Sons, Oslo.

KOutT NIK, D.L. 1984. oo (Euphorbiaceae)—a newly recognized genus in southern

a. S. African J. Bot. 3:262-264.

cn rNIK, D.L. 1987. A taxonomic revision of the Hawaiian species of the genus Chamaesyce
(Euphorbiaceae). Allertonia 4:331—388.

Koutnik, D.L. and M.J. Hurr. 1990. Chamaesyce (Euphorbiaceae). In: W.L. Wagner, D.R.
Herbst, and $.H. Sohmer, Manual of the flowering plants of Hawai'i. Special Publ. ae
Pauahi Bishop Mus. 83, Honolulu. Pp. 602-617.

Lone, R.W. and O. LakeLa. 1976. A flora of tropical Florida. Banyan Books, Miami.

Manaaly, J.K., K. SwARUPANANDAN, and PV. MapHusoopan. 1979. Seed and seedling
morphology of two species of Euphorbia (Euphorbiaceae). Proc. Indian Acad. Sci. 88:87—
94,

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MayeieLb, M. 1991. Euphorbia johnstonii (Euphorbiaceae), a new ir from Tamaulipas,
Mexico, with notes on Euphorbia section Acutae. Sida 14:573-579.

ROEPER, J. ee Enumeratio Euphorbiarum quae in cag et Pannonia gignuntur,
Gottin

ROSENGARTEN or and W.J. HAYDEN. 1983. Stem ontogeny in Chamaesyce hirta (Euphorbiaceae).
ay a et of Science Annual Meeting, Fairfax, Virginia. ABSTRACT: Virginia
dl esvevems|

SIMMONS, a ae W.J. Hayden. 1997. Revision of the cerrado hemicryptophyte Chamaesyce
of Boissier’s eons (Boho Brittonia 49:155—180.

VeH, R.Von. 1928. Beitrag zur Kenntnis der Anisophyllum-Euphorbiaceen und einige
Vergleichende und Entwicklungsgeschichtliche Untersuchungen iiber die Driise des
Cyathiums. Ann. Jard. Bot. Buitenzorg 38:131-162.

WessTer, G.L. 1967. The genera of Euphorbiaceae in the southeastern United States. J.
Arnold Arbor, 48:303—430

WEBSTER, G.L. 1994. Gu snenion of the Euphorbiaceae. Ann. Missouri Bot. Garden.

]

WEBSTER, er W.V. Brown, and B.N. SmitH. 1975. Systematics of photosynthetic car-
bon fixation pathways in Euphorbia. Taxon 24:27-33.

WHEELER, L.C. 1941. Euphorbia subgenus Chamaesyce in Canada and the United States
exclusive of southern Florida. Rhodora 43:97—286

432 SipA 18(2)

W. CHAPMAN, V. CHAPMAN, A. BESSETTE, A. BESSETTE, and D. PENs. 1998.
Wildflowers of New York in Color. (ISBN 0-8 1 15627467, hbk.; ISBN
0-8156-0470-X, pbk.). Syracuse University Press, 1600 Jamesville Avenue,
Syracuse, NY 13244-5160. 315-443-5547; fax 315-443-5545;
<talitz@summon3.syr.edu>. $24.95 (pbk), $59.95 (hbk). 163 pp, 350

color photos.

This book is a popular field picture guide to the wildflowers of New York State, based principally
on Dick Mitchell and Gordon Tucker's The Revised Checklist of New York State Plants. \t is
arranged so that one first must identify the color and structure (symmetry) of the flowers,
then look to habitat (terrestrial vs. aquatic), then to determine if there are leaves, and if
they are present, whether they are basal, alternate, opposite or whorled. The common name,
Latin name and authority, family, oui: season, floral description, plant description,

habitat, and comments are given for each of 350 of the most common wildflowers found
in the state, as well as some of the rare and endangered taxa. The table of contents, visual
glossary, glossary of terms, index to common names, and index to genera and species, makes
finding one’s way around the book extremely easy. The chapter “how to use this guide”
will allow anyone to use the book right away, v with virtually no instruction needed, The
photographs are superb, with excellent depth-of-field and vibrant colors. The high quality,
alkaline paper is also a plus, as is the sturdy, yet flexible cover on the paperback. I ign seen
few other wildflower books of this caliber made for the general public. The authors are to
be congratulated for their fine work and no botanist eaycling through New York State, or
from the Northeast, should be without it.—/ohn J. Pipoly Il

=

Stipa 18(2): 432. 1998

A REVISION OF THE GENUS ARDISTA SUBGENUS
GRAPHARDISIA (MYRSINACEAE)

JOHN J. PIPOLY III

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
jpipoly @brit.org; clusia@latinmail.com

JON M. RICKETSON

Missouri Botanical Garden
> O. Box
St. Louis, MO 63166-0299, U.S.A.

jricketson @lehmann. mobot.org

ABSTRACT

A taxonomic revision of Ardisia subgenus Graphardisia is presented, including an emended
description of the subgenus, a key to the species and subspecies, along with updated de-
scriptions and new illustrations for each taxon. Six taxa are si eee including three
new combinations: Ardisia opegrapha subsp. paquitensis (Lundell) Pipoly & Ricketson, A.
opegrapha subsp. wagneri (Mez) Pipoly & Ricketson, and A. bartlettii subsp. lilacina (Lundell)
Pipoly & Ricketson. Four binomials, Ardisia weberbaueri, A. opegrapha, A. wagner 2 and A.
albovirens, are lectotypified and twen vly rel dto synonymy. Ardisia nigrovirens

acbr. is relegated to synonymy ae A a Meo and both are excluded from

subechus Graphardisia.

RESUMEN

Se presenta una revisién taxonémica del género Ardisia subgénero Graphardisia. Se provée
una descripcién actualizada del subgénero junto con una clave para identificar las especies
y subespecies, descripciones acutalizadas y nuevas ilustraciones para cada taxon. Se reconoce
seis taxa, incluyendo tres combinaciones nuevas: Ardisia opegrapha subsp. paquitensis (Lundell)
Pipoly & Ricketson, A. opegrapha subsp. wagneri (Mez) Pipoly & Ricketson, y A. bartlettii
subsp. /ilacina (Lundell) Pipoly & Ricketson. Se lectotipifican cuatro binomiales: Ardisia
weberbauert, A. opegrapha, A. wagneri y A. albovirens y se relega veintiuno a la sinonimia.
También, Ardisia nigrovirens se relega bajo sinonimia de A. albovirens, y ambas se excluyen
del nee Graphardisia.

INTRODUCTION
The pantropical genus Ardisia Sw. is the largest within the Myrsinaceae,
containing perhaps as many as 500 species (Chen & Pipoly 1996). The
genus has been separated from all others in the Myrsinaceae by the free
filaments and pluriseriate ovules (Mez 1902). While only a few groups within
Ardisia sensu latissimo have been segregated as separate genera in the Paleotropics
Sipa 18(2): 433-472. 1998

434 Sipa 18(2)

(Sadiria Mez (1902), Afrardisia Mez (1902), Tetrardisia Mez. (1902), Hymenandra
A. DC. (1834), Parardisia Nayar & Giri (1986)), there has been an enor-
mous increase in separation of species groups from Ardisia to new genera
in the Neotropics, starting with those who recognize Aublet’s (1775), lcacorea
(against which Ardisia is conserved), followed by Alphonse de Candolle (1841),
Ducke (1930), and finally Cyrus Lundell (1963, 1964, 1981a, 1981b, 1981c,
1981d, 1982). Lundell’s contribution was clearly the most extensive, con-
sisting of the segregation of Amatlania Lundell, Auriculardisia Lundell, Chontalesia
Lundell, Ynckeria Lundell (=Ctenardisia Ducke), Gentlea Lundell, Graphardisia
(Mez) Lundell, /6arrea Lundell, Oerstedianthus Lundell, Synardisia (Mez)
Lundell, Valerioanthus (Mez) Lundell, and Zunilia Lundell. Because the group
is comprised of over 800 names, it will be some time before each species
has been reviewed and the entire group is revised. In the meantime, we suggest
using the key to the Mesoamerican genera we published (Ricketson & Pipoly
1997) to identify specimens to generic level (Ardisia versus Synardisia, Ctenardisia
or Gentlea). For groups within Ardisia, the use of Flora of Guatemala (Lundell
1966a) and Flora of Panama (Lundell 1971) will permit rough identifica-
tion, at least to species groups, until our review of the entire complex is
complete. While most of the taxa described in the remaining segregated
groups are best placed in Ardisia, it appears that some currently placed in
Icacorea and Chontalesia may prove to be problematic. Treatment of those
taxa is now underway (Pipoly & Ricketson, in prep.).

Ardisia subgenus Graphardisia Mez (1902) was described in the Das
Pflanzenreich treatment of the Myrsinaceae and was comprised of only two
species. Ardisia opegrapha Oerst. and A. wagneri Mez. Mez considered sub-
genus Graphardisia allied to Ardisia subg. Synardisia Mez {=Synardisia (Mez)
Lundell] by its imbricate or contorted (but not valvate) corolla lobes and
the included (rather than exserted) stamens. Subgenus Graphardisia was
separated from subg. Synardisia by the rotate (not infundibuliform) corolla,
with lobes connate to only 1/3 (not 3/4) their length. He also considered
Ardisia subg. Graphardisia closely related to A. subg. Ardisia, because of
the often persistent inflorescence bracts and floral bud shape, and contrasted
these two subgenera with A. subg. /cacorea (Aubl.) Mez, known for its very

1conspicuous inflorescence br: acts.

” Lundell (1966b) treated subgenus Graphardisia, but later raised Graphardisia
to generic rank (Lundell 1981c). He (Lundell 198 1c) stated that it was most
closely related i Cas Lundell, because both groups of species
y apical pores. Ocrstedianthus (Lundell
19816) is is comprised of a group including Ardisia nigrescens Oerst., A. tuerckheimii
Donn. Sm., and eight other taxa, formerly placed by Mez (1902) in subge-
nus /cacorea, but segregated from /cacorea by Lundell because of its glan-
dular trichomes of the branchlets, inflorescence and perianth parts. Lundell

=

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 435

(1981c) separated Graphardisia from Oerstedianthus by the following key:
“1. Filaments strictly glabrous; stems and inflorescence glabrous; puncta-

tion of all parts dense and blackish; bracts and bractlets usually folia-
ceous and often persisitent; sepals and cc large, accrescent, usually

blackened, usually ribbed or with dense elevated black glan nds. Graphardisia
1. Filaments pubescent with gland- tipped hairs; stems an ae Dry

glabrous, usuall y puberulent, | hir tellous ora |

and sepals small, not accrescent; sepals not ribbed an Slandsrs iin Oerstedianthus”

Upon examination of the six taxa we recognize in Ardisia subgenus
Graphardisia, 4 taxa (comprising two species in our classification) have ei-
ther stalked glandular papillae or sessile glandular granules on the filaments
near the base, thus invalidating the first part of Lundell’s lead couplet. While
members of the Oerstedianthus species group have anthers very similar to
that of A. subg. Graphardisia, the unique glandular tomentum, and small,
corymbose inflorescences are much different than the panicles of corymbs,
and essentially glabrous plants of subgenus Graphardisia. Our studies indi-
cate that Ardisia subg. Graphardisia is most closely related to A. subg. Icacorea
based on its well-developed panicles with flowers in terminal corymbs; apiculate,
subapically poricidal, concolorous anthers, and style two to three times longer
than the ovary. The two subgenera may be separated by the following key:

1. Branchlets and inflorescence rachises glabrous or rarely, sparsely glandu-
lar-granulose; leaf blades, inflorescence and floral bracts, and perianth
densely and prominently black or red punctate and/or punctate-lineate;
Soe acotens usually clasping as developing fruit Ardisia subg. Graphardisia
ils lepidote; leaf blades rarely
Cnevel prominently punctate -lin eate, the inflorescence
and floral bracts mostly orange, rarely black, punctate, the perianth mostly
orange, rarely black punctate; sepals not accrescent, never clasping the
developing fruit. Ardisia subg. Icacorea

|
u

While preparing a treatment of Ardisia subg. Graphardisia as part of the
Myrsinaceae for Flora Mesoamericana, we treated the entire subgenus so that
it might serve as a precursor to our treatment of the tribe Ardisieae for Flora
Neotropica, that is now in preparation. The revision here presented pro-
vides a nomenclator for Ardisia subg. Graphardisia, that is beyond the for-
mat limitations of the Flora Mesoamericana.

NOTES ON KEYS, TAXONOMIC CONCEPTS, TERMINOLOGY

The keys are artificial and designed to expedite identification of herbarium
specimens. An attempt has been made to emphasize vegetative characters
to increase the keys’ usefulness with sterile material. The numbers appear-
ing before the taxa refer to their respective position in the key; any correla-
tions with phylogenetic relationships are coincidental. Quantitative and

436 Stipa 18(2)

qualitative data presented in keys and descriptions for floral parts and bracts
were taken from organs rehydrated from herbarium specimens by boiling
in water. Measurements from these range from 10% to 15% greater than
those measurements taken directly from dried material. Data regarding stem
diameters, inflorescence rachises, pedicels, leafiand fruit shape and size were
taken from dried herbarium specimens.

Our concept of subspecies follows that of Pipoly (1987) who defined
subspecies as: “groups of populations within a single lineage of ancestor-
descendant populations that show variation by unique combinations of
plesiomorphies, or homoplasic apomorphies, correlated with biogeography
and/or ecology. This rank is primarily used to convey information regard-
ing variation in the life histories of these populations and character state
differences hypothesized to be the result of this variation. The subspecific
rank in no way attempts to predict speciation events.

Morphological terms in this treatment follow Lindley (1848) and Pipoly
(1987, 1992) for the inflorescence, rachis pedicels and floral parts. Description
of leaf morphology follows Hickey (1984), trichome description follows
Theobald et al. (1984) and basic cell and tissue terminology follow Metcalfe
(1984)

TAXONOMIC TREATMENT
Ardisia subgenus Graphardisia Mez in Engl., Pflanzenr. IV. 236 (Heft 9):78.
1902; Lundell, Wrightia 3:192-198. 1966. Graphardisia (Mez) Lundell, Phytologia

48:139. 1981; Lundell, Phytologia 59:429-433. 1986. TYPE: Ardisia opeerapha Oerst.
(LECTOTYPE, here designated).

Mez (1902) included both Ardisia opegrapha as well as A. wagneri in his new subge-
nus. Subsequently, Lundell (1966b) did not designate a lectotype. Therefore, we have
chosen A. opegrapha because it most closely fits the original circumscription of the
subgenus.

Subshrubs to trees. Branchlets mostly terete, glabrous or rarely, glandular-
eranulose. Leaves petiolate; blades membranaceous to subcoriaceous, densely
and prominently black (rarely pellucid-) punctate and punctate-lineate, the
margins entire to crenulate, rarely sharply and irregularly dentate. /nflores-
cence terminal, uni- to tripinnately paniculate, pyramidal to obpyramidal
rarely globose, the ultimate branches corymbose, at times in high anthotactic
spirals and thus appearing umbellate, the rachis often densely and promi-
nently black punctate and punctate-lineate; inflorescence and floral bracts
foliaceous, mostly persistent, resembling the vegetative leaves but acropetally
reduced tn size. aie pink, lavender or purple, densely and prominently
black punctate and punctate-lineate; calyx with sepals free or nearly free,
large, accrescent and clasping fruits at maturity; corolla rotate, the lobes
imbricate in bud, basally short-connate and sparsely to densely yellow glandular-

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 437

granulose within, densely and prominently black punctate-lineate, stamens
inserted at corolla tube base, the filaments basally connate to form an in-
conspicuous tube, the tube free from the corolla tube, the apically free portions
of the filaments glandular-granulose or glandular-papillate, less than half
the length of the anthers, the anthers ovoid-lanceoloid, linear or lanceoloid,
prominently apiculate, dehiscent by subapical pores; ovary globose, the style
lender, equalling the sepals, 2-3 times longer than the ovary, the placenta

apiculate, the ovules pluriseriate, biseriate, or apparently uniseriate (few in
number and in a very high anthotactic spiral). Fruit globose or oblongoid,
densely punctate and punctate-lineate,basally surrounded by persistent, clasping
sepals.

Distribution —A small distinctive subgenus of three species with five subspecies
found from Nicaragua to Bolivia and adjacent Brazil.

Ecology.—Members of the subgenus occur in diverse vegetation types,
including wet and pluvial lowland, premontane, montane, and cloud forests.

The subgenus is defined by: 1) glabrous branchlets and inflorescence rachises;
2) dense and prominently raised black, or rarely, reddish-black, puncta-
tions or punctate-lineations on all leaf and floral parts; 3) sepals often accrescent
and usually clasping the developing fruit; 5) linear-lanceolate, concolorous,
apiculate anthers with subapically poricidal dehiscence; and 6) style 2-3
times longer than the ovary. Species of the subgenus are often used for home
decoration and for use in Christian churches for religious holidays (Pipoly,

pers. obs.).

KEY TO TAXA OF ARDISTIA SUBGENUS GRAPHARDISIA

1. Stoloniferous subshrubs mostly less than 1(—2) m tall; leaf blade mar-
gins sharply and irregularly dentate; corolla tube and filaments yellow
glandular-granulose; Ecuador to Bolivia and adjacent Brazil........... . A. weberbaueri
1. Shrubs to small trees (0.5—)2— Os Le m seat without stolons; leaf blade
m argins entire undulate « or glandula eranulose;
filaments glabrous or sessile to stalked aes Peeillace, Nicaragua
to Colom
2. Sepals So eee oblong, 4.2-8 mm long, apically broadly rounded
to obtuse, hyaline throughout, the margins entire; filaments sessile-
to stalked- glandular-papillate. 2. A. opegrapha
3. Inflorescence obpyramidal; leaf blades oblanceolate or rarely obo-

vate, 3.5-7.5(-8) cm wide, 3 or more times longer than wide.
4. Floral bracts caducous; stamens 5.2-6.5 mm long; filaments
2.5—3 mm long; sepals 5-8 mm long. ...2a. A. opegrapha subsp. opegrapha
4. Floral bracts persistent; stamens 3.8—5 mm long; ee 1.5—
2 mm long; ‘sepals 4,2—5.2 mm long. «-++++ . opegrapha subsp. wagner
3. a globose; leaf blades elliptic to broadly elliptic (7.5-)

aoide2 2—2.5 times longer than

Senne eres reereenenees

Je A. opegrapha subsp. paquitensis

438 SIDA 18(2)
2. Sepals chartaceous, ovate, 1.5—2.5 mm long, apically acute, opaque

except at margin, the margins subentire to erose; filaments glabrous
3. A. bartlettii

5. Sepals 1.5—1.8 mm long; petal lobes 6—6.5 mm long; stamens 3.5—

4.8 mm long; style base tapering; tall wet pOrests. ssaesionseinecbaphtee
a. A. bartlettii subsp. bartlettii
5. Sepals 2—2.5 mm long; petal lobes 7-8 mm stamens 4.5—5.7

mm long; style base stylopodic; strand vegetation and beach for-

ests. 3b. A. bartlettii subsp. lilacina

1. Ardisia weberbaueri Mez, (Figs. 1D, 2), Repert. Spec. Nov. Regni Veg.
3:97. 1906. Graphardisia weberbaueri (Mez) Lundell, Wrightia 7:46. 1982. TYPE:
PERU. JUNIN: Prov. Tarma, near La Merced in Chanchamoyo Valley, without elev.,
Dec 1902 (fl), A. Weberbauer 1809 (HOLOTYPE: B-destr.; LECTOTYPE, here designated: F).

The holotype of Ardisia weberbaueri was destroyed in Berlin (B) in 1943 during WWII.
According to /ndex Herbariorum (Holmgren et al. 1990), other institutions housing
duplicate A. Weberbauer collections are: F MOL, USM and WRSL. Through the
kindness of Carlos Reynel (MOL), Asuncién Cano (USM), and Krzysztof Swierkosz
(WRSL), th rorough searches were conducted at their respective institutions, all fail-

ing to locate isotypes of A. Weberbauer 1809. A fragment of the holotype, including
a mature flower and leaf fragment, was taken from B in 1926, and is available at F (a
“clastotype”—see Pipoly 1983), and is here selected as the lectotype.

Ardisia vigoi Lundell, Wrightia 6:94. 1979. SyN. Nov. sales oe (Lundell) Lundell,
Phytologia 48:140. 1981. Type: PERU. SAN Martin: Prov. Mariscal Caceres, Dtto.
Tocache Nuevo, Rio de la Plata, NE of Tocache, 500-600 m, ; May | 975 (fl, fr), //.
Schunke Vigo 8384 (HOLOTYPE: LL-TEX; tsoryre: MO).

=
~

=
— ~

|

A B Cc D E F
Fic. 1. Comparison of sepal size, shape and areas pe the sae of Ardisia subgen.
Graphardisia. See individual taxon descriptions for details bsp. opegrapha
drawn from G. Webster 16867. B. Ardisia opegrapha ae >. wagnert, re awn from ie D. Chez
24093. C. Ardisia opegrapha subsp. acti drawn fom Araquistain 3007. D. Ardisia
weberbaueri, drawn from R. Vasquez 5924. E. Ardisia bartlettii subsp. bartlettii, drawn from
H. Bartlett & T. Lasser 16720. ¥. Ardisia bartlettii subsp. lilacina, drawn from J. Dwyer 4354.

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 439

Fic. 2. Ardisia weberbaueri Mez. A. Habit, flowering branch. B. Detail of abaxial leaf sur-
face. C. Detail of inflorescence. D. Detail of flower, showing calyx and pistil. E. Detail of
flower, showing corolla and stamens. F. Detail of stamen, showing adaxial and abaxial sur-

faces. G. Fruit. A G drawn from R. Vasquez 5924.

440) Sipa 18(2)

Stoloniferous subshrubs to shrubs, 0.75—1.0(—2.0) m tall. Branchlets slender,
terete, 2-4 mm in diam., glabrous to scattered glandular-granulose. Leaves
alternate; blades membranaceous to chartaceous, oblong to elliptic, 8.2—
19.4 cm long, 1.9-6.6 cm wide apically acuminate, the acumen S—9 mm
long, basally acute to acuminate, decurrent on the petiole, the midrib im-
pressed above, prominently raised below, the secondary veins 22—26 pairs,
obscure above, prominent below, conspicuously punctate and punctate-lineate,
glabrous, the margins irregularly dentate, the teeth 0.4—0.8 mm long, mostly
flat; petioles slender, canaliculate. 0.7-2.1 cm long, glabrous. Inflorescence
terminal, 2.5—7 cm long, 2.5-8 cm wide, usually shorter than the leaves,
the rachis conspicuously punctate and punctate-lineate, glabrous to sparsely
scattered glandular-grandulose; peduncle 2-5 mm long; inflorescence branch
bracts foliaceous, membranaceous, oblong or elliptic, 1.4—2.5 cm long, 0.5—
0.8 cm wide, apically acute to rounded, the midrib impressed above, prominently
raised below, the secondary veins obscure above and below, conspicuously
punctate and punctate-lineate, glabrous, the margins entire to undulate;
petioles 0.8—1.2 mm long, glabrous; inflorescence branches 2—9 mm long,
conspicuously punctate and punctate-lineate, glabrous to scattered glan-
dular-grandulose, bearing terminal corymbs; floral bracts resembling sec-
ondary inflorescence branch bracts, but 7.2-8.1 mm long, 1.7-2.2 mm
wide, sessile or nearly so; pedicels 11.6-13.5 mm long, slender, mostly accescent
in fruit, glabrous to scattered glandular-grandulose. Flowers 5-merous (oc-
casionally with sepals 6-7), membranaceous, 4.5—5.5 mm long; calyx with
sepals free, oblong, 3.2-4.2 mm long, 1.4—1.6 mm wide, apically broadly
obtuse to rounded, densely and prominently punctate and punctate-lineate,
glabrous, except densely glandular-grandulose within basally, the margins
hyaline, subentire, glandular-ciliolate; corolla rotate, 4.3—5.3 mm long, the
tube 0.4—0.5 mm long, densely yellow glandular-grandulose within apically
above staminal tube and below corolla tube and lobe junction, the lobes
ovate, 4.1—4.8 mm long, 2.3-2.5 mm wide, apically rounded to obtuse,
prominently punctate and punctate-lineate, glabrous, the margins entire;
stamens 3.2—3.8 mm long, connate by their bases and adnate to corolla to
form a tube 0.4—0.5 mm long, the free portion 2.8—3.3 mm long, the fila-
ments 1.2—1.4 mm long, 0.5—0.7 mm wide at base, conspicuously punc-
tate, glabrous except scattered sessile glandular-papilllate basally, the an-
thers linear-lanceoloid, 2.0-2.2 mm long, 0.9-1.0 mm wide at base, apically
apiculate, basally sagittate, cent by subapical pores, the connective
prominently black punctate dorsally; pistil obturbinate, the ovary glabrous,
the style 1.9—2.7 mm long, slender, tapering, punctate, glabrous, the ovules
13-14, pluriseriate. /ruitglobose, 6.1-6.8 mm in diam., densely and ca
punctate and punctate-lineate, the style base persistent, glabrou

Distribution.—Ardisia weberbaueri is endemic to the junction the Andes

PIroLy AND RICKETSON, Revision of Ardisia subgenus Graphardisia 44]

and Western Amazonia, from Napo and Pastaza Provinces, Ecuador, southward
through the Departments of Loreto, San Martin, Junin and Madre de Dios,
Peru, to the Departments of Beni, La Paz and Santa Cruz, Bolivia and the
adjacent state of Acre, Brazil. It grows from 180—1,100 m in elevation. This
is the first report of the species for Ecuador.

Ecology and conservation status.—Ardisia weberbaueri is restricted to tall,
lowland and premontane tropical moist and wet forests, where it is an un-
derstory subshrub, growing in loose detritus in sheltered areas near rocks
and tree buttresses. While it is certainly not common, at this time, there
are no data to suggest the species is threatened.

Etymology—Ardisia weberbauert is named in honor of August Weberbauer
(1871-1948), a German botanist and phytogeographer who collected in
Peru from 1901-1905 and again from 1908-1948 (Stafleu & Cowan 1988).
During the 1920s, he was sponsored by the Field Museum of Natural His-
tory (F), under J. Francis MacBride’s Flora of Peru project. Many of Weberbauer’s
collections are type specimens today. Ardisia weberbaueri’s taxonomic syn-
onym, Ardisia vigoi Lundell, was named in honor of José Schunke Vigo,
who had a long collaborative history with the Missouri Botanical Garden
(MO), Field Museum of Natural History (F) and Peruvian Herbaria’s re-
vived Flora of Peru project. José is the son of Carlos Schunke, who was guide
and field assistant to J. Francis MacBride of the Field Museum and G.S.
Bryan of the University of Wisconsin, during their 1923 botanical expedi-
tion to Peru (Dahlgren 1936). Unfortunately, Lundell (1979), confused
José’s second surname (maternal) with his legal (paternal) one, hence the
epithet “vigoi” instead of “schunkei.”

Additional specimen mined. ECUADOR. Napo: Laguna de Yuturi, along both sides
of Rio Yuturi, 00° 36' S, 76° 01' W, 220 m, 26 Feb 1990 (fr), 7. Jaramillo & A. Grijalva
11402 (K); Cantén Orellana, Yasuni National Park, Maxus road and pipeline under con-
struction, km 3 of NPF-Puerto Maxus branch, 04° 01' 00" S, 76° 25' 00" W, 250 m, 10
Jun 1994 (fr), N. Pitman e& M. Aulestia 232 (BRIT, Kk MO). Pastaza: Cantén Pastaza,
“Masaramu’ Oil well, of UNOCAL, 40 km NNE of Montalvo, 00° 44' S, 76° 52' W, 400
m, 1-16 May 1990 (fr), &. Gudifio 355 (BRIT, MO, QCNE). PERU. Huanuco: Prov.
Pachitea, Region Pucallpa, W part, “Siraé Mountains,” and adjacent lowland; 20-24 oe
SE of Puerto Inca, Crest of mountain range going W to E from “Campamento Oro,”

“Campamento Sira” 09° 28' S, 74° 47' W, 600 m, 20 Dec 1987 (fr), B. Wallnéfer 16.
201287 se ih MO, USM, W), 700 m, 8 Apr 1988 (fr), B. Wallnéfer 18-8488 (BRIT,
MO, W). Junin: Prov. Chanchamayo, Chanchamayo, without elev., 22 Oct 1863 (fl), /
an 2234 (EF, MA). Loreto: Prov. Maynas, Allpahuayo, Estacién HAP, without elev., 13
Nov 1984 (fl), R. Vasquez et al. 5924 (MQ); Prov. Maynas, Iquitos, Estacién Experimen-
tal IIAP, Allpahuayo, 21 km al S de Iquitos, 4° 10' S, 73° 30' W, 180 m, 19 Sep 1990
(ster.), J. Pipoly et al. 12168 (MO). Madre de Dios: Prov. Manu, Manu, Parque Nacional
Manu, Cocha Juarez, Rio Manu, 400 m, 3-5 May 1987 (fr), P Nifez et al. 8053 (MO),
San Martin: Prov. Huallaga, Chazute, Rio Huallaga, 260 m, Mar 1935 (fl), G. Klug 3978
(F GH, MOQ); Prov. Mariscal Caceres, Quebrada Huicte, Rio Huallaga, 590 m, 26 May
1964 (fr), /. Schunke V. 6449 (MO); Dtto. Tocache Nuevo, Quebrada de Huaqguisha, right

442 Sipa 18(2)
bank of Rio Huallaga, without elev., 17 May 1970 (fr), J. Schunke V. 3983 (F), 8 Jun 1970
(fr), J. Schunke V. 4039 (F); Puerto Pizana, right bank of Rio Huallaga, lout elev., 3
Sep 1971 (fr), J. Schunke V. 4983 (F, MO); Quebrada de Cachiyacu de Lopuna, Progreso
Hwy., 500-850 m, 21 Jul 1974 (fl, fr) ee Schunke V. 7647 (MO); Quebrada Paraiso, 5 km
below Tocache, 400 m, 27 Nov 1974 (fl, fr), 7. Schunke V. 8133 (MO). BOLIVIA. Beni:
Prov. Ballividn, lower slopes of Serranfa Pilén Lajas, 14.3 km N of the bridge over the Rio
Quiquibey, 15° 19'S, 67° 03' W, 700 m, 10 Jun 1985 (fr), 7. Solomon 13940 (MO); Misién
‘4tima, S of San Borja, near Rfo Maniquf, on afluent of Rfo Chimané, behind the ce
240 m, 20 May 1988 (fr), S. Beck 16323 (MO); E slopes, 21 km from Yucumo, 15° 1
67° 04' O, 1,035 m, 17 May 1989 (fr), D. Smith et al. 13188 (MO). La Paz: Prov. ee
Yungas, Alto Madidi, ridge top ca. 7 km NE of camp, 13° 35' S, 68° 46' W, 300 m, 27
May 1990 (fr), A. Gentry & S. Estensoro 70712 (MO); Alto Beni, trail from bridge toward
San Antonio, 570 m, 3 Jan 1988 (fr), R. Seidel & M. Schulte 2316 (MO); Prov. Sud Yungas,
Alto Beni, Sapecho Concession, Yungas, 15° 30'S, 67° 20' W, 550 m, 3 Apr 1989 (fr),
Seidel et al. 2671 (MO), 3 Feb 1990, R. Seidel & V. Baptista 2824 (MO). Santa Cruz:
Prov. Ichilo, Parque Nacional Amboré, 3 km SW of Estancia San Rafael de Amboré, 17
km by air, SSE of Buena Vista, forest near Rio Surutu, 17° 36'S, 63° 37' W, 350 m, 29 Jul
1987 (fr), M. Nee et al. 35424 (MO, TEX); ca. 15 km (SE) up the Rio Pitasama from the
Rio Suruti, 17° 44' S, 63° 40' W, 700 m, 28 Aug 1985 (fr), /. Solomon 14104 (MO).
RAZIL. Acre: Rio es Seringal, S40 Francisco, without elev., Oct 1911 (fl), E. Ule 9641
(K).

Ardisia weberbaueri appears to be most closely related to A. bartlettii, based
on its sepals that are opaque except along the margin, relatively small inflo-
rescences with early caducous floral bracts, and non-entire margined leaf
blades. However, Ardisia weberbaueri may be easily separated from A. bartlettii
by its diminutive, stoloniferous habit, sharp and irregular teeth of the leaf
blades, and the glandular-granulose filaments.

Detailed study of the lectotype of A. weberbaueri and the type of A. vigor
revealed no quantitative or qualitative difference between the two respec-
tive type specimens.

2. Ardisia opegrapha Oerst.

Shrubs or small trees (0.6—)2—10 m tall. Branchlets (2-)3—5(—7) mm in
diam., glabrous throughout. Leaves alternate or pseudoverticillate; blades
membranaceous, elliptic, broadly elliptic, oblanceolate, or obovate, 10—30
cm long, 3.5—-14.5 cm wide, apically acuminate, base acute or attenuate,
decurrent on the petiole; midvein impressed above, prominently raised below,
the secondary veins prominulous above, inconspicuous below, densely and
conspicuously punctate and punctate-lineate, glabrous, the margins entire
or irregular; petioles canaliculate and usually marginate, 0.2—-3.5 cm long,

glabrous. /nflorescence terminal, a bipinnate, radially symmetrical (globose)
or obpyramidal panicle of corymbs, 2-12 cm long, 2-15 cm ae usually
shorter than the leaves; peduncle obsolete to 3.5 cm long, conspicuously
punctate and punctate- lineate, glabrous, inflorescence branch bracts mem-
branaceous, widely ovate, 7.2-26 mm long, 1.4—8 mm wide, apically broadly

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 443

rounded, densely and prominently punctate and punctate-lineate, glabrous,
caducous or persistent in fruit, the margins entire; floral bracts as in sec-
ondary branch bracts except oblong, 3.7—12 mm long, 0.8—5.5 mm wide;
pedicels 6-23 mm long, slender, conspicuously punctate and punctate lineate,
sparsely glandular-glandulose. Flowers 5-merous, membranaceous, 7.5—10
mm long; calyx with sepals free, linear, oblong or broadly elliptic, 4.2-8
mm long, 2.2-3.1 mm wide; apically rounded, emarginate, densely and
prominently punctate and punctate-lineate, very sparsely glandular- -granulose
without, densely glandular-granulose at base within, the margins entire,
sparsely glandular-ciliolate or not; corolla rotate, 6.5—9.5 mm long, the tube
0.3-0.7 mm densely yellow glandular-granulose within apically above staminal
tube as well as below corolla tube and lobe junction, the lobes ovate to el-
liptic, 6.5—9 mm long, 3.8-5.7 mm wide, apically rounded, densely and
prominently punctate and punctate-lineate, glabrous, the margins entire;
stamens 3.8—6.5 mm long, the staminal tube 0.3—0.8 mm long, the fila-
ments 1-3 mm long, 0.5—0.8 mm wide, epunctate, scattered glandular-
papillate, the anthers linear-lanceoloid, 2.3-3.6 mm long, 0.9-1.1 mm wide
at base, apiculate, basally sagittate, concolorous, dehiscent by subapical pores,
the connective inconspicously punctate dorsally; pistil obturbinate, the ovary
glabrous, the style 3.5-5 mm long, slender, punctate and punctate-lineate,
glabrous, the ovules 11-13, pluriseriate. Fruit globose, 4.2-8 mm in diam.,
densely and prominently punctate, glabrous.

Ardisia opegrapha, as here circumscribed, includes three newly recognized
subspecies for which combinations are made herewith.

2a. Ardisia opegrapha Oersted subsp. opegrapha (Figs. 1A, 3). Ardisia of
Oerst., Vidensk. Meddel. Dansk Naturhist. Foren Kjabenhavn 1861: 126. 1861
Graphardisia opegrapha (Oerst.) Lundell, Phytologia 48:140. 1981. TYPE: COS’ TA.
RICA. ALAJUELA: Aguacate, without elev., Nov 1847 (fl), A. Ocersted 29A (LECTO-
TYPE, here designated: C, (F Neg. # 22954); ISOLECTOTYPE: F)

—

In the Botanical Museum, University of Copenhagen Herbarium (C), there are two
different collections that were cited in the protologue by Oersted, two duplicates of
A.S. Oersted 29 from Jaris, and one of A. Oers ted 29A from Aguacate, of which only
A. Oersted 29A is duplicated in the Field Museum wie History (F). The collec-
tion from Jaris was made in November 1846, and that from Aguacate in November
1847. In Oersted’s original description he states “Crescit in kee costaricencibus
Aguacate et Jaris (1500—2000’), ubi florentem et fructificantem Novembri legi,” without
further specifying the location. Mez (1902) and subsequent authors have not desig-
nated a lectotype. We here select the collection from Aguacate at C as the lectotype
because it contains the greatest number of dissections in the original fragment packet,
indicating it was a more critical element in the description of the species.
eg ae Mast., Gard. Chron., n.s., 8:680, fig. 132. 1877. SYN. Nov. non Bot. Mag

er. 3, vol. 34]:¢. 6357. 1878,. Graphardisia oliveri (Mast.) Lundell, Bivilbe
ye 285. 1983. Type: COSTA RICA. PRov. UNKNOWN: From Messrs. Veitch & Sons
Nursery, Jul 1876 (fl), A. Endres s.n. (HOLOTYPE: K, (LL-TEX Neg. # 71-146); Isoryres:

444 Sipa 18(2)

Fic. 3. Ardisia pegrapha Oersted subsp. opegrapha. A. Habit, flowering branch. B. Detail

of abaxial leaf surfac . Detail of inflorescence. D, Detail of stamen, showing adaxial

surface. E. Detail of stamen, showing laterial surface. F Detail of stamen, showing abaxial
surface. G. Fruit. A, B drawn from lectotype. C—F drawn from G. Webster 16516. G drawn
from R. Mendoza 26.

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 445

W 3-sheets).

Ardisia setbertii Stand]., Ann. Missouri Bot. Gard. 24:198. 1937. SYN. Nov. Graphardisia

seibertii (Standl.) Lundell, Phytologia 48:140. 1981. Type: PANAMA. Coc e: EI Valle
ntén and vicinity, 500-700 m, 23-27 Jul 1935 (fl, fr), RJ. Serbert 456 (HOLO-
TYPE: F, (F Neg. # 68250); isotypes: A, K, MO (LL-TEX Neg. # 71-102)).

Ardisia skutchii C. V. Morton, J. Wash. Acad. Sci. 27:309. 1937. SYN. Nov. Graphardisia
skutchti (Morton) Lundell, Phytologia 59:432. 1986. Type: COSTA RICA. SAN José:
onan of El General, 1,070 m, Jun 1936 (fl), A. Skutch 2660 (HOLOTYPE: US (CM

g. # 49641, LL-TEX Neg. # 71-104, US Neg. # 2386); isotype: MO (LL-TEX
Noe 71-104)).

Ae subcoriacea Lundell, Wrightia 3:193. 1966. SYN. Nov. Graphardisia subcoriacea
(Lundell) Lundell, Phytologia 48:140. 1981. Type: PANAMA. Coc te: El Valle de
Anton, vicinity of La Mesa, ca. 1,000 m, 22 Jun 1941 (fl), PH. Allen 2571 (HOLO-
TYPE: US, (LL-TEX Neg. # 71-111); isotypes: A, F (F Neg. # 68253)).

Ardisia picturata Lundell, Wrightia 4:164. 1971. SYN. Nov. Graphardisia ee (Lundell)
Lundell, Phytologia 48: oe 1981. Type: PANAMA. PaNaMA: Cerro Campana area,
above Su Lin Motel, ca. 3,000 ft [904 m], 11 Aug 1967 (fr), /. D. Dwyer & J.H.
Kirkbride Jr. 7818 (HOLOTYPE: LL-TEX n.v.,; isoTyres: K n.v., LL-TEX, MO, NY, UC
n.v., cee

Graphardisia coib cundell, Reece 59:429. 1986. SYN. Nov. Ardisia ee Sens
Lundell, Phyrologi 61:63. 1986, nomen invalidum. Type: PANAMA. Veracuas: Playa
Rosario, N tip of Coiba Island, without elev., 26 Aug 1970 ( - R. Foster 1600 (HO-
LOTYPE: LL- ‘TEX; isoTyPes: DUKE n.v., F (F Neg. # 68357), MO).

Graphare s Lundell, Phytologia, 59:429. 1986. SyN. Nov. Ardisia esquipulasana
handel Phyoloss 61:63. 1986, nomen novum invalidum, non Ardisia nicaraguensis
Oerst., Vidensk. Meddel. Dansk Naturhist. Foren Kjobenhavn 1861133. 1861. Type:
NICARAGUA. Zetaya: Montafia Esquipulas, 130 m, 22 Nov 1951 (fr), PJ. Shank
& A. Molina R. 4719 (HoLorype: F (F Neg. # 68356); isorype: US).

Graphardisia obtusata Lundell, Phytologia 59:430. 1986. Syn. Nov. Ardisia obtusata (Lundell)
Lundell, Phytologia 61:66. 1986, nomen invalidum. Type: PANAMA. Darlin: Rio
Tuquesa, at middle Tuquesa Mining Company camp called Charco Peje, ca. 250 m,
8 le rae (fr), S. Mori 7034 (HOLOTYPE: LL-TEX; isorype: MO).

ay ylla Lundell, japan 59:430. 1986. SYN. Nov. Ardisia riomonteana
ests see 61:66. 1986, nomen novum invalidum, non Ardisia oxyphylla
Wall., Numer. List 2291. 1830. Type: PANAMA. CuiriQut: 2.5 km from Questa Piedra
along Rio Monte Road, at stream, without elev., 27 Jun 1977 (fl), 7. P Folsom 3975
(HOLOTYPE: LL-TEX; isotypes: CR n.v., MEXU n.v., MO, NY, PMA n.v.

Graphardisia arte Denes Eeyore 59:432. 1986. SYN. Nov. Ardisia ustupoana
(Lundell) Lundell, P logia 61:68. 1986, nomen novum invalidum. Type: PANAMA.
SAN Bias: through clean on mainland in front of Ustupo, overhanging river,

without elev., 9 Nov 1975 (fr), WG. D'Arcy 9535 (HoLoTyPE: LL-TEX; isotypes: BRIT,
MO)

See

Graphardisia murphyae Lundell, Phytologia 63:77. 1987. SYN. Nov. Ardisia murphyae
(Lundell) Lundell, Phytologia 63:463. 1987. Type: COSTA RICA. PUNTARENAS: N
of La Lucha road to Progresso, 1,200 m, 23 May 1984 (fl), H. Murphy 1248 (HOLO-
Type: DUKE, n.v., LL-TEX fragment of holotype).

Graphardisia serenoana [seranoana] Lundell, Phytologia 59:431. 1986. SYN. Nov. Ardisia
seranoana (Lundell) Lundell, Phytologia 61:66. 1986, nomen invalidum. Tyre: PANAMA.
CHIRIQUI: road from Volcan to Rio Sereno [Serano], road that turns E 7.2 km from
Rio Serano, 3.2 km along side road, without elev., 29 Jun 1977 (fl), /.P Folsom 4029
(HOLOTYPE: LL-TEX; Isoryrpes: MEXU n.v., MO, PMA n.v.

446 Stipa 18(2)

Shrubs or small trees (A-)2—10 m tall, 2—5 cm in diam. Branchlets 3-5
mm in diam. Leaves with blades elliptic to lanceolate, broadly elliptic or
oblanceolate, 12-25 cm long, 3.5—7.5 cm wide; petioles canaliculate, usu-
ally marginate, 0.2-1.5 cm long. /nflorescence obpyramidal, bipinnately
paniculate, 2-12 cm long, 4-15 cm wide; peduncle 0.5—1.5 cm long; in-
florescence branch bracts widely ovate, 16-18 mm long, 14-16 mm wide,
caducous; floral bracts 7-12 mm long, 3—5.5 mm wide, oblong, caducous;
pedicels 10-23 mm long. Flowers 9-10 mm long; sepals linear, oblong to
broadly elliptic, 5-8 mm long, 2.2-2.5 mm wide; corolla 8—9.5 mm long,
the tube 0.4—0.6 mm long, the lobes oblong to broadly elliptic 7.6-8.9
mm long, 4.5—5.5 mm wide; stamens 5.2—6.5 mm long, the staminal tube
0.4-0.7 mm long, the filaments 2.5—3 mm long, 0.6—-0.8 mm wide, the
anthers 2.7—3 mm long, 0.9-1 mm wide at base; ovary with style 4-5 mm
long, the ovules 12-many. Fruit S—8 mm in diam.

Distribution.—Nicaragua (Jinotega, Matagalpa, Zelaya, Rio San Juan),
Costa Rica (Alajuela, Puntarenas, San José), Panama (Chiriqui, Veraguas,
Herrera, Los Santos, Coclé, Panama, San Blas, Darién) and Colombia (Choco);
sea level—2,500 m elevation. It is notable that the subspecies principally
inhabits areas facing the Pacific coast of Central and adjacent South America.

Ecology and conservation status.—Ardisia opegrapha subsp. opegrapha in-
habits a wide range of habitats, including beach thickets; gallery forests in
deciduous woodlands and dry oak forests; moist, wet, lower montane, montane,
cloud and elfin forests. Because it is mostly found at forest margins, and
along watercourses, subsp. opegrapha may be found in primary and disturbed
or remnant sites.

Etymology.—The epithet ‘opegrapha’ comes from the Greek, ‘opsis, re-
sembling, and graphe, meaning drawing, picture or writing. This refers to
the densely and prominently punctate and punctate-lineate vegetative and
floral parts.

Additional specimens examined. NICARAGUA. Jinotega: Cerro San Pedro, Comarca
Kilambé, 13° 36' N, 85° 38-39' W, 820 m, 21 Jul 1980 (fl), 7. — 192 uae Matagalpa:
Las Brisas, 15 km W of Waslala, road to El Tuma, 13° 15' N, 85° 28' W, 250-300 m, 23
Dec 1982 (fr), P Moreno ee (MQ). Rio San Juan: pueae. Aires, | aoe al N of Caserfo
Sabalo, along Rfo Sdbalo, 11° 02' N, 84° 28' W, 70 m, 10 Sep 1985 (fr), 2 Moreno 26297
(MO); aa Rio — ruz and Cafio Santa Crucita, La Palma, 11° 02—04' N, 84°
24—26' W, 40-60 m, 30 Nov—2 Dec 1984 (fr), W. Stevens et al. 23470 (MO). Zelaya: El
Achiote, 11° 47' N, 84° 26' W, 200 m, 25 Aug 1982 (fr), M. Araquistain 3130 (MO); San
Antonio, 200 m, 29 Aug 1982 (fr), A. ge 27 (MO); along Rio : | Zapote, W of Nueva

, 11° 42'N, 84° 26' W, 130 m, 21 Aug 1983 (fr), /. Miller & J. Sandino 1257
(MO): 4. 4 km N of base camp, base Bele 3.6 km SE of Cerro oT a, Rio Kama, Rio
Escondido, 12° 05—15' N, 83° 45'—84° 20' W, sea level-65 m, 16 Mar 1966 (fr), G. Proc-
tor et al, 2 hoot (NY, L a ‘ EX); 5 km SE de La Providencia, along Cano Chiquito, we
out elev., 21 Aug 1983 (fr), /. Sandino 4534 (MO); Comarca de ELC abo, 40—45 km
of ae 10-100 m, - tan 1970 (fr), E Seymour 3783 (MO); Esquipulas and nee

—

PIPOLy AND RICKETSON, Revision of Ardisia subgenus Graphardisia 447

Rio Alemén drainage, 150 m, 27-29 Nov 1951 (fr), P Shank & A. Molina R. 4783 (F, LL-
TEX, MO). COSTA RICA. Alajuela: toward center of Cerro de San Isidro, San Ramén,
1,150 m, 10 Jul 1925 (fl), A. Brenes 4313 (F, LL-TEX); between Santiago and San José de
San Ramon, without elev., 17 Oct 1928 (fr), A. Brenes 6349 (F); San Miguel de San Ramon,
without elev., 21 Jul 1934 (fl), A. Brenes 19242 (F); Rio Grande, San Isidro, San Ram6n-Alajuela,
1,000 m, 10 Jun 1982 (fr), A. Carvajal 269 (LL-TEX, MQ); 3 km E of San Ramén, 1,025
m, 22 Jun 1969 (fl), R. Lent 1765 (F); Los Angeles de Las Gania, 1,100 m, 21 Dic 1936
(fl), E Solis R. 495 (LL-TEX, MO). Puntarenas: Cantén Coto Brus, Las Cruces Botanical
Garden, Cordillera de Talamanca, 08° 47' 10" N, 82° 58' 30" W, 1,200—1,300 m, 30 Jun
1995 (fr), L. Angulo 388 (BRIT, MO); “Hort. Eisgmb, Clt. Lauche,” without elev., 1895
(fl), C. Beck s.n. (W); Cerro Pando, ridges above the Rfo Cotén and the Rio Negro, near
La Lucha, Rfo Cotén, 08° 55' N, 82° 45' W, 1,000-1,800 m, 19-21 Feb ce (fr), K.
Barringer & L. Diego G. 1652 (F); Pucet Quepos, along stream in shady woods on W side
of National Park, 16 mi E of town, without elev., 28 Feb 1982 (fr), 7, Churchill Mee (F);

foothills of the Cordillera de Talamanca, directly N of Las Alturas, 08° 57' N, 82° 50' W,
1,400—1,500 m, 28 Aug 1983 (fr), G. Davidse 24159 (MO); Fila El Tigre, SE of Las Alturas,
08° 56'N, 82° 51' W, 1,350—1,450 m, 29 Aug 1983 (fr), G. Davidse 24176 (MO); Foot-
hills of Cordillera de Talamanca, forested valley of the Rio Cotén between Sitio Cotén
(Cotoncito), and Sitio Tablas, 08° 57' N, 82° 46' W, 1,500-1,600 m, 2 Sep 1983 (fl), G.

Davidse 24428 (MO); Parque Boltvar, without elev., 10 Jul 1945 (fl), /- reed 4147
(F); La Tigra-Las Mellizas, 1,280 m, Aug 1983 (fl, fr), L. Gomez et al. 21982 (LL-TEX,

MO); Cantén Coto Brus, NE slopes of Fila de Cal, between San Vito de Coto Brus and
Ciudad Neily, 08° 41' N, 82° 56' 50" W, 500-620 m, 12 Jul 1985 (fr), M@. Grayum & B.
Hammel 5638 (MO); Alturas de Coto Brus, ca. 20 km NE of San Vito, 1-3 km S of Las
Alturas, 08° 56' N, 82° 50' W, 1,300-1,400 m, 13 Jul 1985 (fl), B. Hammel & M. Grayum
14201 (MO); Las Cruces Botanical Garden, San Vito, jungle trail, 08° 47’ 08" N, 82° 57'
29" W, 1,250 m, 4 Jul 1994 (fl), W Kress & T. Prinzie 4572 (MO, US); Amistad Biosphere
Reserve, near Las Alturas de Cotén Biological Station, 08° 57' 00.3" N, 82° 49' 56.8" W,
1,455-2,100 m, 8 Jul 1994 (fl), W Kress & T. Flores 4266 (MO, US); 1,475—-1,750 m, 8
Jul 1994 (fl), Wo Kress & T. Prinzie 4582 (BRIT, MO, US); above La Tigra, Talamanca
Range, near Panamanian border, 1,600—1,800 m, 20 Mar 1982 (fr), D. Hazlett 5064 (F);
between Sitio Tigra and Sitio Las Tablas picnic area, near Cerro Chivo, S of Cotén river,
1,600-1,800 m, 17 Jul 1983 (fl), D. Hazlett 5229 (F); beside Rio Bella Vista near Las
Alturas, 08° 56' N, 82° 51' W, 1,300 m, 8 Jul 1972 (fl), R. Lent 2693 (EK MO); Cafias
Gordas, 1,100 m, Feb 1897 (fr), A. Pittier 11190 (LL-TEX); 1 mi S of San Vito de Java,

3,500 ft [1,067 m], 18 Aug 1967 (fr), 2 Raven 21889 (F, MO); Piedra Blanca de Villa

Colén, 1,200 m, 17 Abr 1935 (fl), M@. Valerio 1268 (F); 7.5 km NE of Sabalito, 08° 51' N,

82° 53' W, 1,021 m, 7 Jul 1977 (fl), G. Webster 21876 Hie Finca Loma Linda, SW of
Cafias Gordas, 08° 44’ N, 82° 56' W, 3,600 ft [1,097 m], 17 Jul 1977 (fl), G. Webster
21937 (F); 4 km S of Las Alturas, 1,400 m, 15 Jul 1977 (fl), R. Wilbur et al. 22713 (MO).

San José: forest along Quebrada de los Micos, ca. 8.5 km by road, W of Ciudad Colén,
09° 55' 50" N, 83° 17' 50” W, 600 m, 16 Jan 1986 (fr), M. Grayum & P. Sleeper 6103
(MO); Zona Protectora La Cangre} a, along Rio Negro, ca. 1.5 km E of Santa Rosa de
Puriscal, 09° 42' N, 84° 23' 05" W, 320 m, 14 May 1987 (fl), 4. Grayum et al. 8325
(MO); between Pedernal and Candelaria, 900 m, 29 May 1966 (fl), A. Jiménez 3966 (F,
MO); road from Santiago to Parrita, 600 m, 24 Sep 1967 (fr), Rk. Lent 1321 (F, NY); Zona
Protectora La Cangreja, Santa Rosa de Puriscal, en las margenes del Rio Negro, 09° 42

24" N, 84° 23' 35” W, 400 m, 10 Aug 1992 (fr), 7, Morales 318 (BRIT, MO); “Cultivé,
Jardin Monte Alegre 4 San José”, without elev., without date (fl), A. Pittier 2298 (BR);
vicinity of El General, 1,190 m, Nov 1936 (fl, fr), A. Skutch 2890 (GH, MO), 670 m, Jun
1939 (fl), A. Skutch 4371 (A, FE, K, LL-TEX, MO); Cantén Turrubares, flank of Cerro

448 Stipa 18(2)

Pel6n, Zona Protectora Cerros de Turrubares, 09° 48' 50" N, 84° 28' 00" W, 1,500 m, 7
Nov 1990 (fr), 2. Zuniga et al. 392 (BRIT, MO). PANAMA. Chiriqui: Cerro Colorado,
mining road 15.6 mi above bridge over Rio San Félix, 1,330 m, 21 Nov 1979 (fr), 77 An-
tonio 2573 (LL-TEX, MO); Cerro Colorado, 8.5 km from Escopeta, along stream, 800—
0 m, 17 Aug 1977 (fr), /, Folsom 4925 (LL-TEX, MO). Coclé: vicinity of El Valle,
800—1,000 m, 22 Dec 1936 (fl), P Allen 72 (LL-TEX, MO); vicinity of EI Valle, 800—
1,000 m, 5 Sep 1938 (f], fr), P Allen 786 (LL-TEX, MO); N rim of El Valle, without elev.,
4 Jun 1939 (fl), 2 Allen & A. Alston 1846 (MO); El Valle de Antén, 750 m, 2 Jul 1941 (fl),
P Allen 2577 (A, LL-TEX, MO), 1,000 m, 16 Jun 1946 (fl), P Allen 3535 (BM, MQ); El
Valle de Anton, N hills, without elev., 29 Jun 1946 (fl), P Allen 3561 (F-3-sheets, G, LL-
TEX -2 sheets, MO, MU); El Valle, without elev., 20 Jun 1966 (fl) Ko Blum et al. 2382
(MO); 5 km W of El Valle on dirt road to top of grassy ridge, 3,000 ft [914 m], 26 Feb
1978 (fl), B. Hammel 1764 (LL-TEX, MQ); hills NE of El Valle de Anton, 2,000 ft [610
mJ], 27-29 May 1967 (fl), W Lewis et al. 1803 (LL-TEX, MO); El Valle, on mountain
slope near hotel, without elev., 30 Nov 1975 (fl), R. Mendoza 26 (MO); ca.15-20 km NE
of La Pintada towards Todbre, largest peak in vicinty, 600—1,000 ft [183-305 m], 15 Feb
1981 (fl), K. Sytsma & Wo D’Arcy 3610 (MQ); border Coclé-Panama Prov., N slopes of
Cerro Guacamayo, SE of El Valle, 08° 36' N, 80° 07' W, 1,800 ft [549 m], 3 Jul 1971 (fl),
G. Webster 16867 (MO); between Las Margaritas and El Valle, without elev., 15 Jul-8 Aug
1938 (fl), R. Woodson et al. 1239 (A, LL-TEX, MO), 1746 (MO). Herrera: hill above Chepo
de Las Minas, 700 m, 19 Dec 1977 (fr), /. Folsom et al. 6991 (LL-TEX, MO); W of Las
Minas, on Montosa de Chepo, vicinity of Chepo, 07° 42' N, 80° 51’ W, 900 m, 20 May
1987 (fl), G. McPherson 10933 (MO, PMA). Los Santos: Rio Guanico Valley, 07° 18' N,
80° 30° W, 600 m, 25 May 1986 (fl), G. McPherson 9248 (BRIT, LL-TEX, MO, PMA);
above Guanico River, on hills W of river, 07° 20' N, 80° 30' W, 550-650 m, 5 Jan 1989
(fr), G. McPherson 13513 (BRIT, MO, PMA). Panama: Camino de Plantacién, 09° 04’
N, 79° | 39' W, 80 m, 1 Jun 1995 (ster.), S. Aguilar & A. Castillo 2116 (F); Summit of
Cerro Campana, 800-1,000 m, 1 Sep 1940 (fl), 2 Allen 2226 (F, LL-TEX); Cerro Campana,
trail Campana to Chica, 600-800 m, 10 Aug 1941 (fl), P Allen 2661 (LL-TEX); Interpre-
tation Trail, 1 km E of INRENARE Forest Ranger Camp, 8° 40' N, 79° 55' W, 800—900
m, 2 Aug 1989 (fl), M. Correa et al. 5139 (BRIT, MO, PMA); Cerro Campana above Su
Lin eee without elev., 25 May 1971 (fl), 7 Croat 14746 (MO); Cerro Campana at
turnoff to FSU cabin, without slew. 20 Jul 1974 (fl), T. Croat 25189 (MO); Cerro Campana,
2,500 ft [762 m], 17 Aug 1982 (fr), We D'Arcy & C. Hamilton 14976 (LL-TEX, MO);
erro Campana, upper slopes in National Park, 207 m, 13 May 1980 (fl), D. LeDoux
2%6 30 (MO); Cerro Campana, beyond Su-Lin, 2,700—3,000 ft [823-914 m], 8 Sep 1966
(ster.), /. Duke 8650A (MO); Cerro Azul, 1 mi below Goofy Lake, 2,000 ft [610 m], 21
Jul 1972 (fl), A. Gentry & J. Dwyer 5512 (F, MO);Cerro Campana, slopes S of radio tower,
2,500 ft [762 m], 1 Jul 1978 (fl), B. Hammel 3761 (LL-TEX, MO); between peaks of
Cerro Trinidad, saddle on SE slope, without elev., 5 May 1968 (fl), /. Kirkbride & J. Duke
1642 (MQ); Cerro Campana, without elev., 10 Dec 1967 (fl), W Lewis et al. 3038 (LL-
TEX, MO); Altos de Campana, 285 m fromMotel Su Lin, 951 m, 25 Jun 1977 (fl), R&
Méndez 45 (MO), without elev., 25 Jun 1977 (fl), R. Méndez 47 (MO), 3,045 ft [928 ml],
3 Dic 1977 (fr), Rk. Méndez 80 (F, MO); Cerro Campana, 2,800 ft [853 m], 29 Aug 1965
(fl), S. McDaniel 6912 (MO); W of Inter-American Hwy. near Capira, Cerro Campana
08° 40' N, 79° 50' W, 900 m, 11 Nov 1985 (fl, fr), G. McPherson 7458 (BRIT, F, LL.
TEX, MEXU, MO, PMA, US); Parque Nacional Altos de Campana, Interpretation Trail,
1 km E of forest INRENARE forest ranger cabin, Tree No. $2158, 08° 40 N, 79° 55' W,
800-900 m, 8 Jul 1988 (fl), AZ. Ruiz et al. 165 (BRIT, PMA), 30 Sep 1988 (fr), M4. Ruiz er
al. 210 (BRIT, PMA), 2 Aug 1989 (fl), M1. Rutz et al. 5139 (BRIT, PMA), 23 Aug 1990
(fl), M1. Ruiz et al. 7320 (BRIT, PMA), 23 May 1991 (fl), MZ. Ruiz et al. 7800 (BRIT,

WS

—

—

—

PIPOLY AND RICKETSON, Revision of Ardisia s1 us Graphardisia 449

PMA), 13 Jun 1991 M. Ruiz et al. 7865 (BRIT, PMA); Goofy Lake, SW facing slope, 500
m, 5 Jul 1976 (fl), G. Sullivan 58 (MO); Cerro Campana, 08° 40' N, 79° 55' W, 2,700-
2,800 ft [823-853 m], 11-12 Jun 1971 (fl), G. Webster & G. Breckon 16516 (MO, TEX).
eraguas: Vicinity Santa Fé, forested slopes of Cerro Tute, 2,500 ft [762 m], 25 Mar 1947
(fl), 2 Allen 4404 (F, G, MOQ); vicinity of Escuela Agricultura Alto Piedra, near Santa Fé,
along trail to top of Cerro Tute, 3,600—4,000 ft [1,097—1,219 m], 29 Jun 1980 (fl), 7
Antonio 4961 (LL-TEX, MQ); Islas Contreras, Isla Uva, 08° 48' N, 81° 45' W, 50 m, 18
Jul 1984 (fr), H. Churchill 5687 (LL-TEX, MO); along road on Pacific slope 1-3 km above
Escuela Agricola Alto Piedra, 700-800 m, 26 Jul 1974 (fl), Z Croat 25995 (LL-TEX, MO);
between Santa Fé and Escuela Agricola Alto de Piedras, without elev., 29 Aug 1974 (fr), 7!
Croat 27350 (LL-TEX, MO); 0.2 mi beyond fork in road at Escuela Agricola Alto de Pie-
dra on road to Rio Calovebora, 750 m, 3 Apr 1976 (fl, fr), 7 Croat & J. Folsom 33865
(LL-TEX, MQ); above Santa Fé beyond Escuela Agricola Inter-Americana, 1.8 mi beyond
fork in road on Pacific slope, on side of Cerro Tute, without ae , 5 Apr 1976 (fr), 7 Croat
34205 (LL-TEX, MO); along road between Santa Fé and Rfo Calovebora, 1.8 mi beyond
Escuela Agricola Alto de Piedra, 735 m, 5 Apr 1976 (fr), 77 mee Folsom 34267 (MO);vicinity
of Santa Fé, along dirt road past ee Circlo Alto de Piedra, formerly Escuela Agricola
Alto de Piedra, to Rio Luis, along Rio Primero Brazo, first stream below school, on Atlan-
tic Coast, 08° 33' N, 81° 08' W, 490 m, 28 Jun 1987 (fl), 7’ Croat 66873 (BRIT, LL-TEX,
MO, PMA); Parque Nacional, Cerro Tute, vicinity of Santa Fé, along road between Alto
Piedra, on slopes of Cerro Tute, 800—1,030 m, 15 Jul 1994 (fl), 7) Croat & G. Zhu 76926
(MO); 7 km NW on road to Santa Fé, without elev., 5 Dec 1975 (fr), W. D'Arcy 10274
(MQ); SW side of Cerro Tute, La Cuchilla, 2,500 m, 10 Sep 1982 (fl), W. D'Arcy 15019
(MQ); Isla de Coiba, without elev., 18 Aug 1961 (fl, fr), 7, Dwyer 1612 (F); along beach of
Isla de Coiba, without elev., 28 Jul 1962 (fr), (, Dwyer 2363 (MEXU, NY); traditional
campsite, 1 km past Ag School, forest slope to the rear, road from Santa Fé, 1,000—1,200
m, 5 Feb 1977 (fr), Folsom & L. Collins 1613 (MO); shoulder of Cerro Tuti, without
elev., 25 May 1977 (fl), /. poe a al. 3375 (MO); Cerro Tute ridge up from former Escuela
Agricola, Santa Fé, 08° 35' N, 81° 05' W, 800-1,100 m, 15 Jul 1983 (fl), C. Hamilton &
ie Krager 3973: (1-1 EX, or trail on ridge to summit of Cerro Tute, Cordillera de
Tute, 1 km past Escuela Agricola Altos de Piedras, W of Santa Fé, 08° 36' N, 81° 06' W,
750-950 m, 15 Dec 1981 (fl), S. Knapp & K. Sytsma 2498 (LL-TEX, MO), (fr), S. Knapp
eK. Sytsma 2518 (LL-TEX, MO); above Escuela Agricola Alto de Piedra, just W of Santa
Fé, 3,200 ft [975 m], 7 Jun 1973 (fl), , Luteyn 4012 (LL-TEX); N of Santa Fé, on prop-
erty of Escuela Agricola Alto de Piedra, woods near cattle shelter at Girasol, 16 Oct 1974
(fl), S. Mori & J. Kallunki 2520 (LL-TEX, MO); Cerro Tute, ca. 10 km NW of Santa Fé,
on lower slopes, 750—1,000 m, 19 Jun 1975 (fl), S. Mori 6738 (LL-TEX, MO), 2 Aug
1975 (fl), S. Mori et al. 7545 (LL-TEX, MO); Cerro Tute, along ridge-trial towards sum-
mit, 08° 30' N, 81° 07' W, 1,000—1,250 m, 21 Mar 1987 (fr), G. McPherson 10744 (MO);
vicinity of Cerro Tute-Arizona, along trail to summit beyond Altos de Piedra, above Santa
Fé, 08° 30' N, 81° 10’ W, 850-1,100 m, 27 Jul 1988 (fl), G. McPherson 12797 (BRIT, F,
LL-TEX, MO, PMA); along trail to summit of Cerro Tute, ca. 3 km above Escuela Agricultura
Alto Piedra near Santa Fé, 2600-2800 ft [792-853 m], 4 Jan 1981 (fr), Ki Sytsma & T.
Antonio 2997 (BRIT, MO). COLOMBIA. Chocé: Quebrada Changamé to the mouth of
the Rio Jurubida, 06° 05' N, 77° 10' W, sea level-100 m, 5 May 1990 (fl), C. Barbosa
6412 (BRIT, MO); trail between Curiche and Alto de Curiche, 10—1,000 m, 22 Jun 1968
(fl, fr), /. Duke 15799 (MO, OS)
It is notable that Ardisia coibana (Lundell) Lundell, A. esquipulasana Lundell,
A. obtusata (Lundell) Lundell, A. riomonteana Lundell, and A. ustupoana
(Lundell) Lundell, are all invalid names because Lundell (1986) made the

oO.

450 Stipa 18(2)

transfers “to obviate the incentive of workers to undertake such unneces-
sary reductions [of /cacorea (Aubl.) Mez, Auriculardisia Lundell, Gentlea
Lundell, Graphardisia (Mez) Lundell, Ocrstedianthus Lundell, barrea Lundell,
Amatlania Lundell, and Zunilia Lundell/ to subgenera.” Therefore, according

34.1, we feel that the names are invalidly published because both
clauses, “(a) when it is not accepted by the author in the original publica-
tion, and (b) when it is merely proposed in anticipation of the future ac-
ceptance of the group concerned, or of a particular circumscription, posi-
tion or rank of the group,” are clearly fulfilled. However, this is not the
case for Ardisia murphyae (Lundell) Lundell because the transfer (Lundell
1987) was effected with no commentary on the part of the author. We have
not validated Lundell’s combinations in Ardisia here because we consider
these names taxonomic synonyms and to do so would require that we rec-
ognize them as distinct taxa.

Ardisia opegrapha subsp. opegrapha exhibits great quantitative variation
among relative size of its organs within individuals and among populations
within its somewhat restricted geographic range. Repeated sampling from
one individual at Parque Nacional Altos de Campana, Panama (M/. Ruiz et
al. 165, 210, 5139, 7320, 7800, 7865) over a three year period revealed
that relative size of leaves to inflorescence is highly variable even within the
same individual. Given that the inflorescences are terminal, the branches
are pseudoverticillate and growth is rythmic, Ardisia opegrapha exhibits Scarrone's
Model (Hallé et al. 1978). Repeated collections would amount to a prun-
ing experiment and while the tree grew from 1.5 m tall when first collected
in July of 1988 (VM. Ruiz et al. 165) and reached a height of three meters in
June of 1991 (M. Ruiz et al. 7865), some of the radical size difference among
the leaves on the specimens may be consequent to reiteration phenomena
previously discussed by Pipoly (1992, 1998). Unfortunately, no data is available
to determine whether successive specimens were taken from normal sym-
podial (by substitution) branches of the tree, or from reiterative (bayonet
type) shoots as a direct result of pruning effected during specimen collec-
tion. Morphological variation among populations is much greater than that
observed within individuals, and this plasticity has led to overdescription,

—

because species circumscription has been reliant on relative size of inflores-
cences to leaves, bracts to pedicels, relative crowding of the flowers in the
corymbs, length of floral parts, and plant height. The rationale for relegat-
ing 12 species to synonymy is explained below; relative terms (larger, smaller,
thinner, and thicker) refer to comparisons with populations with features
of the type of Ardisia opegrapha.

Ardisia oliveri was collected from populations whose inflorescences are
larger than average, large relative to the leaves, and whose flowers were in
full anthesis. Ardiséa seibertii also has inflorescences that are large relative

PipoLy AND RICKETSON, Revision of Ardisia subgenus Graphardisia 451

to the size of the leaves; the inflorescence branches bear pedicels in high
anthotactic spirals, giving the impression that the flowers are in umbels rather
than true corymbs. The type of Ardisia skutchii is notable only for its sepals
that are slightly shorter and wider than the average, short petioles and flowers
white with only a slight pink tinge. Ardisia subcoriacea was described be-
cause it has relatively smaller, subcoriaceous leaves, petioles not marginate
to the base, smaller flowers, elliptic sepals with hyaline margins and short
stamens. Graphardisia coibana, whose type is a fruiting collection, was separated
by its thick leaves with abruptly caudate-acuminate apices and long-attenuate
bases. However, both of these features lie well within the range of quanti-
tative variation for the subspecies.

Ardisia picturata’s type represents populations with extremely thin leaves,
but otherwise identical to “typical” subspecies opegrapha. It is interesting
to note that the type (Dwyer e& Kirkbride 7818) is a specimen that amply
demonstrates the range of variation in size and shape of the leaves within
the taxon; the LL-TEX isotype has small nearly elliptic leaves, while the
isotypes at MO and NY have leaves nearly twice as large as the LL-TEX
sheet, and are elliptic to widely oblanceolate (essentially obovate). Clearly,
there is no practical way to separate the A. picturata morphotype. Other
taxa segregated principally because of thin leaves include Graphardisia
nicaraguensis and G. obtusata, both of which are otherwise notable for their
oblanceolate and oblong leaves, the former with obovate or elliptic sepals
and the latter with narrowly obtuse sepal apices. Recognition of these vari-
ous taxa amounts to describing plants different in overall aspect rather than

unique characters.

Other examples of minor variations used to separate species include the
sparsely ciliolate sepals found on the type of Graphardisia ustupoana, and
the pellucid rather than black punctate leaves found on the type of Graphardisia
murphyae. In addition, Graphardisia oxyphylla has reddish instead of black
punctate-lineate sepals, relatively thick petioles and flower size at the smaller
end of the continuum for the subspecies. The description of Graphardisia
seranoana was based primarily on its subsessile anthers, but reexamination
of the type reveals that the measurements were taken from very young flo-

ral buds. The duplicate collection at MO contained more mature buds,
and open floral remnants with stamens well within the size range of varia-
tion found in subsp. opegrapha. Vherefore the mistake was owing to the
holotype specimen’s condition rather than to a biological characteristic.

Ardisia opegrapha is most closely related to A. bartlettii based on its yel-
low glandular-granulose corolla tube, but can easily be separated from it
by the membranaceous, oblong apically broadly rounded to obtuse sepals
with entire margins, and sessile- to stalked glandular-papillate filaments.

Among the subspecies of Ardisia opegrapha, subspecies opegrapha is most

452 SipA 18(2)

easily recognized by a combination of the obpyramidal inflorescence, ca-
ducous floral bracts and floral parts larger than the other two subspecies. It
appears to be more closely related to subsp. wagneri than it is to subsp. paquitensis
because of the obpyramidal inflorescence shape and leaves 3 or more times
longer than wide.

2b. Ardisia opegrapha Oerst. subsp. wagneri (Mez) Pipoly & Ricketson,
comb. et stat. nov. (Figs. 1B, 4). Ardisia wagneri Mez, Pflanzenr. IV. 236 (Heft
9):79. 1902. Graphardisia wagneri (Mez) Lundell, Phytologia 48:141. 1981. TYPE:
PANAMA. CHIRIQUE: Volcan de Chiriqui [Volcan de nee without elev., without
date (fr), M. Wagner 623 (LECTOTYPE, ate designated: GOET).

Mez (1902) listed collections from Volcan Baru, including M. ‘i 623 and F Lehmann
sm., and cited herbaria at GOET, M, and W, without reference as to which specimen
was deposited where. Searches of the collections at M and W failed to discover any
material of either M/. Wagner 623 or E Lehmann s.n. A misidentified collection of
Lehmann s.n. was located at W, bearing a determination by Mez in 1901 as Ardisia
opegrapha. Therefore, we assume it aoe not fit Mez’s original concept of A, wagnert.

"he Wagner collection at GOET most closely fits the description of A. wagneri and
we therefore designate it as the lectotype.

Ardisia bracteolata Lundell, Wrightia 6:65. 1979. SYN. Nov. Graphardisia bracteolata (Lundell)
Lundell, Phytologia 48:140. 1981. Type: NICARAGUA. Zexaya: 6.3 km S of bridge
at Colonia Yolaina on road to Colonia Man. antiales of Nueva Guinea, primary tall,
evergreen forest on steep areas above stream, 200-300 m, 13-14 Feb 1978 (fr), PC.
Vincelli 252 (HOLOTYPE: MQ; ISOTYPE:

Ardisia zelayensis Lundell, Wrightia 6:95. 1979. SYN. NOV. Graphardisia zelayensis (Lundell)
Lundell, Phytologia 48:141. 1981. Type: NICARAGUA. Zetaya: near Rio Okwamwas,

m E of Rosita, without elev., 15 Jun 1978 (fl), D. Neill 4478 (HoLtotype: MO (F

Neg. # 55686)).

Graphardisia hyalina Lundell, Wrightia 7:273. 1984. Syn. Nov. Ardisia neohyalina Lundell,
Phytologia 61:65. 1986, nomen novum invalidum, non Ardisia hyalina Lundell, Wrightia
3:99. 1964. Type: COSTA RICA. ALAJUELA: Finca Los Ensayos, ca. 11 mi NW of
Zarcero, primary forest and ieee ca. 850 m, 15 Aug 1977 (fl), 7. Croat 43565
(HOLOTYPE: LL-TEX; isorype: MO).

Graphardisia purpurea Lundell, Phytologia 59:431. 1986. SYN. Nov. Ardisia ebingeri Lundell,

rytologia 61:63. 1986, nomen novum invalidum, non Ardisia purpurea Reinw. in

Blume, Bijdr. 13:684. 1826, Tyee: PANAMA. Cuiriqut: Boquete, 6 mi N of Concepcién,
second growth forest, without elev., 4 Aug 1960 (fr), LE. Ebinger 751 ee
US; IisoType: MO)

Shrubs to small trees (0.5—)2—6 m tall, 2-5 cm in diam. Branchlets 2-4
mm in diam. Leaves with blades elliptic or oblanceolate 10-23 cm long,
4—7.5(—8) cm wide; petioles canaliculate, 0.7—2 cm long. /nflorescence
obpyramidal, a pinnate panicle of corymbs, 2-8.5 cm long, 4-15 cm wide;
peduncle obsolete to 3.5 cm long; inflorescence branch bracts numerous,
oblong, 10-26 mm long, 3.5-8 mm wide, persistent; floral bracts linear to
oblong, 6—9.2 mm long, 1—3.1 mm wide, persistent; pedicels 6-23 mm
long. Flowers 8—9.2 mm long; sepals oblong or widely oblong, 4.2—5.2 mm

SS
-

joy

<o

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 453

ors

2
7
D

Fic. 4. Ardisia opegrapha Oersted subsp. wagneri (Mez) Pipoly & Ricketson. A. Habit,
flowering branch. B. Detail of abaxial leaf surface. C. Detail of inflorescence. D. Detail of
flower, showing corolla and stamens. E. Detail of stamen, showing adaxial and abaxial surf:
Fruit. A, B, drawn from lectotype. C, F drawn from P Moreno & W. Robleto 20745. D-
E, drawn from F Araya et al. 306.

454 S1pA 18(2)

long, 2.2-3.1 mm wide; corolla 7.1-9 mm long, the tube 0.3—-0.5 mm,
the lobes ovate, 7.0-8.5 mm long, 3.8—4.5 mm wide; stamens 3.8—5 mm
long, the staminal tube 0.3—0.6 mm long, the filaments (0.5—)1.5—2 mm
long, 0.5—0.7 mm wide, the anthers 2.3-3.2 mm long, 0.9—1.1 mm wide
at base; ovary with style 3.5—5 mm long; ovules 12-13, pluriseriate. Fruit
4,2—5.5 mm in diam.

Distribution.—Central America, primarily in areas with drainage toward
the Caribbean, subsp. paquitensis is found from Nicaragua (Jinotega, Matagalpa,
Chontales, Zelaya and Rio San Juan) southward through Costa Rica (Guanacaste,
Alajuela, Heredia, Puntarenas and Limén) to Panama (Chiriqui and Bocas
del Toro) from sea level—1450 m elevation.

Ecology and conservation status.—Primary, remnant and secondary for-
ests in lower montane, premontane, evergreen tropical wet forests and cloud
forests.

Etymology— The specific epithet commemorates Moritz Friedrich Wagner
(1813-1887), Bavarian botanist and zoologist, who traveled to Panama and
Ecuador between 1857—1860

Additional specimens examined. NICARAGUA. Chontales: Santo Domingo, 500—1000
m, 13 Jan 1970 (fl, fr), aes 3346 (MQ); Cerro Oluma, near top of Cordillera Amerisque
12° 18'N, 85° 24' W, 840 m, 3 Jan 1984 (fr), A. Gentry et al. 43897 (MO); Cerro Olen:
ca. 3 km SW de miler a. 12° 18' N, 85° 20' W, 700-740 m, 3 Jan 1984 (fl), A.
Grijalva et al. 3380B (MQ); Cerro Las Nubes, El Tamagas and Loma San Gregorio, ca. 2
km N of Santo Domingo, 600 m, 20 Jan1984 (fr), A. Grijalva & D. Rios 3493 (MO); 4
km NNW of Cuapa, ridgetops and summits of Cerro Oluma, 12° 18' N, 85° 23' 30" W,

700-775 m, 23 Sep 1983 (fl), M4. Nee 28425 (MO). Jinotega: Comarca Santa Cruz, el
Calvario, al SW del Cerro Kilambé, 13° 34' N, 85° 40' W, 900-1,000 m, 27 Mar 1981
(fr), P Moreno 7730 (MQ); Las Brisas, Comarca Kilambé, 13° 35—36' N, 85° 39' W, 930
m, Jul 1980 (fl), 7. Sandino 160 (MO); Cerro San Pedro, Comarca Kilambé, 13° 30’
N, 85° 38-39' W, 820 m, 21 Jul 1980 (fl), /. Sandino 186 (MO). Matagalpa: 78 from
Minas along Matagalpa-Siuna Hwy, La Gloria, 13° 15' N, 85° 35' W, 600 m, 13 Sep

1982 (fr), P Moreno 17227 (MO); El Comején, 1 km W a Hwy. to Waslala, 13° oe N,

85° 34’ W, 600 m, 23 Feb 1983 (fr), P Moreno 20597 (MO). Rio San Juan: Along Rio
Sabalo, 11° 03' N, 84° 28' W, 40 m, 07—08 Jul 1984 (fl), P Moreno & W. Robleto 25981
(MQ); La Lupe near Rio Santa Cruz, 11° 08' N, 84° 22' W, without elev., 7 Oct 1990 (fr),
J. Salick 7860 (MO). Zelaya: Nueva Guinea, 11° 46' N, 84° 26' 30" W, 200 m, 11 Ago

1982 (fl), M. Araquistain 3007 (MO); Nueva Guinea, Colonia Yolaina, 11° 40' N re
22' 30" W, without elev., 13 Ago 1982 (fl), M. Araquistain 3087 (MQ); El ehioie. 1°
47'N, 84° 26' W, 200 m, 25 Ago 1982 (fl), M. Araquistain 3132 (MO); Cerro El Naranjo,
ca. 15 km SW of Wani, without elev., 14 Sep 1982 (fl), A. Grijalva e» P Moreno 1134
(MO), (fr), A. Grijalva e~ P Moreno 1153 (MO); Rio Punta Gorda, Atlanta, La Richard,
Loma San Jorge, 11° 31' N, 84° 04’ W, 150-160 m, 13 Nov 1981 (fr), 2 Moreno & J.
eis 13039 (MQ); Rio Punta Gorda, Atlanta, al SE de La Richard, 11° 32' N, 84° 05'

0 m, 14 Nov 1981 (fr), 2 Moreno & J. Sandino 13100 (MO); Comarca Waslala, 6.5
are - SE de Waslala, 13° 16' N, 85° 24' W, 520-560 m, 14 Sep 1982 (fr), P Moreno

17273 (MO); El Naranjo, a 50 km de Siuna, along Matagalpa-Siuna Hwy., 10 km al W
de la cuesta El Guayabo, 13° 34' N, 85° 10' W, 300 m, 14 Sep 1982 (fr), P Moreno 17312
(MQ); La Posolera, 5 km al W de Waslala, carretera El] Tuma a Waslala, 13° 17' N, 85° 24'

-—

PIpoty AND RICKETSON, Revision of Ardisia subgenus Graphardisia 455

W, 700 m, 22 Dic 1982 (fr), P Moreno 19119 (MO); El Gudsimo, camino a El Dos, NE
de Siuna, 13° 48' N, 84° 39' W, 360-380 m, 25 Feb 1983 (fr), P Moreno e& W. Robleto
20745 (MO); Mpio. de Siuna, Wanf, without elev., 27 Ago 1982 (fr), F Ortiz 49 (MQ);
Mpio. de Siuna, Comarca Santa Rosa, without elev., 6 Sep 1982 (fr), E Ortiz 112 (MO);
Mpio. de Siuna, Comarca Danlf, 100-130 m, 31 Sep 1982 (fl), F Ortiz 201 (MO); Mpio.
de Siuna, Waspado, 100-120 m, 6 Oct 1982 (fr), F Ortiz 281 (MO); Mpio. de Siuna, El
Torno, 120 m, 10 Oct 1982 (fr), F Ortiz 324 (MO); Sector Mina Nueva América, 13° 45'
N, 84° 30' W, 500 m, 22 Sep 1984 (fr), F Ortéz 2130 (MO); Bonanza, on ground of
Neptune Mining Co, 14° 02' N, 84° 35' W, 350-450 m, 26 Feb 1979 (fr), /. Pipoly 3516
(MO); Laguna Santa Rosita on He fro BBO nas to El Salto Grande, 14° 03' N, 84° 37'
W, 140-160 m, 27 Feb 1979 (fl), /. Pipoly 3670 (MO); Finca Waylawas, 13° 30) N, 84°
48' W, 80 m, 12 Mar 1979 J. Pipoly 4420 (MO); Risco de Oro, 40 m, 22 Mar 1979
(fl), J. Pipoly 5010 (MO); Estacién Experimental El Recreo, 12° 09' N, 84° 17' W, 15 m,
10 Jan 1985 (fr), D. Rios 232 (MO); 1 km S de Colonia Verdun, en Nia Guinea, 11°
38' N, 84° 26' W, 200-250 m, 17 Jul 1982 (fl), 7. Sandino 3260 (MO); Cerro El Escobin,
4 km de Colonia Serrano, 11° 33-34' N, 84° 21-22' W, 120-130 m, 30 Jul 1982 (fl), /.
Sandino 3369 (MO); 1 km de Colonia Serrano, Rio Serrano, 11° 34' N, 84° 21-22' W,
70-80 m, 31 Jul 1982 (fl), 7. Sendino 3427 (MO); Toro Bayo and Esquipulas, drainage of
Rios Jicaro and Esquipulas, 130 m, 20 Nov 1951 (fr), 2 Shank e A. Molina R. 4610 (F);
Esquipulas Mountains, 130 m, 22 Nov 1951 (fr), P Shank & A. Molina R. 4696 (F); Esquipulas
Mountains and Aleman, Rio Aleman Drainage Basin, 150 m, 27-29 Nov 1951 (fr), 2
Shank & A. Molina R. 4766 (F), (ster.), P Shank & A. Molina R. 4850 (F); Rios Punta
Gorda, Aleman, and Zapote Drainage Basins, 30 m, 5 Dic 1951 (fl), P Shank eA. Molina
R. 4966 (F); Estacién Experimental El Recreo, ca. 1.5 km SE of the station, 100 m, 10
Ene 1985 (fr), D. Soza et al. 378 (MO). along new road from Rfo Blanco to Rio Copalar,
ca. 31 km E of Rio Blanco, 12° 50-55' N, 85° 00—05' W, 200-400 m, 13 Feb 1979 (fl),
W. Stevens 12153 (MO, NY); along road from Bonanza to Constancia, 13° 58'-14° 01’
N, 84° 37-40' W, 160-360 m, 21 Feb 1979 (fl), W. Stevens 12493 (MQ); along road be-
tween El Empalme and Limbaika, ca. 1.5 km SE of Palmera, 13° 35' N, 84° 20' W, 60 m,
24 Feb 1979 (fr), W Stevens 12836 (MO); along Rfo Waspuk ca. 1 km upstream from
confluence of Rio Pis-Pis, 14° 15' N, 84° 36 W, 75-100 m, 1 Mar 1979 (fr), W. Stevens
13083 (MO). Without Department: 1867 (fl), B. ae s.n. (BM), COSTA RICA. Alajuela:
Cantén San Ramon, R.F. San Ramon, Cordillera de Tilaran, trail to Volcdn Muerto, 10°
12' 55" N, 84° 36' 25" W, 1,100—-1,200 m, 11 May 1993 (fl), & Araya et al. 306 (BRIT,
MO, NY); Reserva Biolédgica Monteverde Rio Pefias Blancas, Parcela de Manuel Rojas,
850 m, 8 Aug 1988 (fl), £. Bello 291 (MO); Canton Upala, ca. 3 km NNE of Bijagua
along new road to Upala, 10° 45' N, 85° 03' W, 450 m, 7-8 Nov 1975 (fr), W! Burger &
R. Baker 9847 (F); E slopes of Volcan Miravalles, W of Bijagua, near the Rio Zapote, 10°
44'N, 85° 05' W, 600 m, 11-12 Feb 1982 (fr), We Burger et al. 11684 (BM, F, LL-TEX);
along road between Canas (Guancaste) and Upala, near Rio Zapote, 1.8—2.7 km S of Rio
Canalete, 100 m, 25 Jun 1976 (fl), 77 Croat 36349 (BRIT, MO); along Hwy. 15, between
Naranjo and Aguas Zarcas, 8 km NE of Quesada, 600 m, 3 Feb 1979 (fr), 7) Croat 46928
MO); without further locality, without elev., without date (fl), E. Friedrichsthal 1296 (W);
Canton Upala, RN. Guanacaste, Cordillera de Guanacaste Estacién San Ramon, La Campana,
Dos Rifos, 10° 52' 50" N, 85° 24' 05" W, 550 m, 23 Mar 1994 (fl), 77 Garcia 203 (BRIT,
MO); Laguna de Rfo Cuarto, Sarapiqui, 400 m, 5 Nov 1984 (fr), 7. Gémez-Laurito 10215
(F); along creek in Florencia San Carlos, 250 m, 30 Jun 1985 (fl), W Haber et al. 1770
(MQ); Cantén San Ramon, Valley of Rio La Esperanza, 4 km SW of La Tigra, Finca Araya
Ledezma, Bosque Eterno de Los Nifios, 10° 18' N, 84° 37' W, 500-600 m, 3 Jul 1992 (fl),
W. Haber et al. 11260 (MO); Cantén Upala, Dos Rios, 5 km S of Brasilia, right bank of
Rio Pizote to Minga House, 10° 55' N, 85° 20' W, 500 m, 31 Oct 1987 (fr), G. Herrera

—

456 Sipa 18(2)

1060 (MO); Canton San Carlos, along Rio Sey near La Marina, Llanura de San Carlos,
550 m, 21 Feb 1966 (fr), A. Molina et al. 17708 (F); Parque Nacional Rincén de la Vieja
‘olonia Blanca, farm in Quebrada Rancho ae 10° 46' 55" N, 85° 15' 10" W, 500
m, 14 Jul 1991 (fl), G. Rivera 1452 (MO); S slope of Volcan Arenal, above Rio Agua Caliente,
without elev., 21 Feb 1989 (fr), G. Russell et al. 870 (MO, US); 9.8 km N of Rio Naranjo
on road to Upala, local name for area is El Macho, 440 m, 8 Nov 1975 (fr), /. Utley & K.
Utley 3273 (F); ca. 2 km S of the town of Canalete along road to Upala, 100 m, 12 Nov
1975 (fr), J. Utley & K. Utley 3305 (F); Cantén Aguas Zarcas, Atlantic rain forest area on
Hacienda la Marina, Rio San Rafael, 450-500 m, 8 Feb 1965 (fr), L. Williams et ul 29120
(F). Guanacaste: Canton Tilaran, bosque entre La Laguna del Arenal y el Alto de La Carpintera,
700 m, 26 Jun 1930 (fl), A. Brenes 12653 (F); Z.P. Tenorio, Cordillera de Guanacaste,
Estacion Tenorio, 10° 36' 40" N, 84° 59' 45" W, 1,050 m, 10 Aug 1992 (fl), G. Carballo
436 (MO); Rancho Harold, region da Volcan Cacao, 700—1,200 m, 30 Jul 1986 (f), /.
Chacon e& A. Chacén 2153 (MO); La Cruz de Abangares, 1,400 m, 15 Jul 1985 (fl), W
Haber & E. Bello C. 1995 (MO), W. Haber 2000 (MO), 2079 (MO); 2 km SW of La
Cruz, on J. Wolfe Farm, 1,400 m, 28 Aug 1985 (fl), Wo Haber 2439 (MO), 2440 (LL-
TEX), 2441 (MO), 2442 (MO); La Cruz de Abangares, 1,360 m, 4 Sep 1985 (fl),
Haber & J. Wolfe 2513 (BRIT, LL-TEX, MO), 1,400 m, 10 Sep 1985 (fl), We Haber & E.
Bello C. 2667 (BRIT, LL-TEX, MO), 2691 (LL-TEX), 2693 (MO), 24 Sep 1985 (fl), W
Haber & E. Bello C. 2888 (LL-TEX, MO); oe es Rio Chiquito Zana ones,
Finca de Campos, Vertiente Atlantica, 10° 25' N 3' W, 1,100 m, 7 Jun 1987 (fl), W
Haber ec E. Bello C. 7439 (MO), 7454 (MO, ae a. Liberia, Parque Rincén de La
Vieja, del Mirador siguiendo la Fila al Volcdn Santa Marfa, 10° 46' N, 85° 49' W, 1,100—
1,300 m, 27 Nov 1987 (fl, fr), G. Herrera ie (MO); Parque Nacional Guanacaste Estacién
Picilla, Finca La Pasmompa, 11° 02' N, 85° 24' W, 300 m, 17 Jun 1989 (fl), 7 INBIO 53
(BRIT, MO); Silencio- Vilaran, without na 31 May 1932 (fl), W. Kupper 1529 (M); Cantén
Liberia, Parque Nacional Guanacaste, Cordillera de Guanacaste, Estacién Cacao, trail to
Fran’s House, 10° 55' 43" N, 85° 28' 10" W, 1,100 m, 8 Feb 1995 = M. Lobo 25 (BRIT,
MO); Sendero Nacho y La Esperanza, 10° 59' 26" N, 85° 25' 40" W, 700 m, 22 Jul 1991
(fl), P Rios 385 (MO); Hacienda Tenorio, without elev., 17 Feb 1956 (fl, fr), B. Schubert
1086 (A); ca. 3 km N of Rio Naranjo, near the continental divide and ca. 100 m S of the
Guanacaste—Alajuela border, 10° 42' N, 85° 04' W, 500 m, 5 Jan 1975 (fl), J. Zaylor 18094
(F). Heredia: Carrillo Estacién Magsasay, Sarapiqui, 10° 24' 18" N, 84° 03' 30" W, 200
m, 5 Jul 1990 (fl), D. Acevedo et al. 128 (MO); 5 Jul 1990 (fl), G. Carballo 15] (BRIT,
MO); N of Puerto Viejo, 12 km to ferry, over ferry, 6 km along road, 100 m, 3 Feb 1983
(fr), NV. Garwood et al. 961 (BM); Cantén Sarapiqutf, Parque Nacional Braulio Carrillo,
Estacién Carrillo, Quebrada Sandijuela, 500 m, 25 Jul 1984 (fl), L. Gomez et al. 22966 (F,
LL-TEX, MO); Finca L a Selva, the OTS field station on the Rio Puerto Viejo just E of its
junction with the Rio Sarapiqui, along West River Road, near point, without elev., 16 Aug
1979 (fl), M. Grayum 2403 (MO); edge of trail on point, 100 m, 8 Apr 1982 (fl), B. Hammel
11619 (F), 100 m, 23 Apr 1982 (fl), B. Hammel 11825 (F, LL-TEX); Cantén Sarapiqui,
Finca La Selva, Puerto Viejo, forest along Quebrada Leonél, 10° 26' N, 84° 01' W, with-
out elev., 11 Jul 1973 (fl), G. Hartshorn 1254 (BRIT, F. MO); La Selva, ca. 1,650 m along
Holdridge Trail, at creek crossing, tributary of Quebrada El Pantana, 100 m, 15 Jul 1984
(fl), B. Jacobs 2871 (MO); La Selva, 775 m line, West River Road, 100 m, 22 Oct 1982
(fr), 7’ McDowell 562 (MQ); E of 1120 West River Road, between Rio Puerto Viejo and
Hunters Cocoa grove, 100 m, 1 Dec 1982 (fr), 77 McDowell 1007 (MO); Finca La Selva,
ca. 4 km SE of Puerto Viejo, 10° 28' N, 84° 00' W, 50 m, 21 Jul 1976 (fl, fr), 7. Solomon
5328 (F, MO). Limén: Cantén Pococi, R.N.ES. Barra del Colorado, Llanura de Tortuguero,
Sardinas, 15-20 m, 12 Dec 1992 (fl, fr), F Araya 147 (MO); La Concepcién, Llanuras de
Santa Clara, 250 m, Apr 1896 (fl), 7, Donnell Smith 6677 (LL-TEX); Vicinity of U.S. Department

—

=

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 457

of Agriculture Rubber Experiment Station, Los Diamantes, on Rio St. Clara, 1.6 km E of
Guapiles, 200 m, 9 Jul 1949 (fl), R. Holm e H. Ilris 315 (G, GH); Cantén Powei, La
Granja, Finca Vieja, 260 m, 26 Jul 1936 (fl), F Solis R. 416 (FE, MO). Puntarenas: Monteverde,
Altos del Rio San Luis, Divisién al Rio Lagarto, Santa Elena, 10° 16' N, 84° 50' W, 1,000
m, 18 Jul 1989 (fl), &. Bello C. 1032 (MO); Reserva Bioldgica Monteverde, Altos de Santa
Elena, 10° 19' N, 84° 49' W, 1, 400 m, 6 Aug 1991 (fl), E. Bello C. 2929 (INB, MO);
Monteverde, lower community, 1,350-1,400 m, 1 Mar 1985 (fr), W. Haber 1427 (MO),
1428 (LL-TEX); Monteverde, Pacific slope forest, 1,450 m, 8 Aug 1985 (fl), W Haber
2187 (MO), 2188 (LL-TEX), 2189 (MO), 2190, 2191, 2192, 2193, 2194, 2195, 21%,
2197, 2198, 2199 (MO); Monteverde community, 1,450 m,18 Aug 1985 (fl), W. Haber
2276 (MO), 2315, 2316 (LL-TEX); 2317 (MO), 2318, 2319, 2320, 2321, 2322, 2323,
2324, 2325, 2326, 2327, 2329, 2331 (MO); Monteverde community, Pacific slope, 10°
20' N, 84° 50' W, 1,400 m, 9 Jun 1986 (fl), W Haber & E. Bello C. 5031, 5039, hom
5063 (MO), 16 Sep 1986 (fl), W Haber & E. Bello C. 5641 (MO). Province unknow
without date, (fl), E. Friedrichsthal s.n. (W), (fl), E Lehmann s.n. (W). PANAMA. ihe
del Toro: Water Valley, Vicinity of Chiriquf Lagoon, without elev., 9 Sep 1940 (fl), H. von
Wedel 661 (LL-TEX, MO), 23 Sep 1940 (fl), H. von Wedel 932 (MO), 9 Oct 1940 (fl), H.
von Wedel 1109 (MO), 5 Nov 1940 (fr), H. von Wedel 1536 (LL-TEX, MO), 26 Nov 1940
(fr), H. von Wedel 1769 (MO). Chiriqui: Without further locality, without elev., 1858
(fl), Md. Wagner 246 (M (F Neg. # 20056)); Vicinity of San Bartolomé, Peninsula de Burica,
sea level-50 m, 28 Jul-01 Aug 1940 (fl), R. Woodson & R. Schery 906 (LL-TEX, MO),

As with the other two subspecies of Ardisia opegrapha, marked but con-
tinuous quantitative variation in relative sizes of parts and persistence of
floral bracts has caused much overdescription. The type of Ardisia bracteolata,
a fruiting collection, is notable only for its thin, narrowly oblong sepals
that are at times emarginate apically, often varying within the same speci-
men. Ardisia zelayensis was described owing to its relatively large sepals with
obtuse apices, and the leaves with acute apices and cuneate bases. Popula-
tions represented by the type of Graphardisia hyalina vary from all others
of the subspecies because their anthers have more rounded bases and more
narrow apices. The diminutive size of the inflorescence, its rachis and pedicels,
together with the extremely slender pedicels of populations from which the
type of Graphardisia purpurea was collected probably led to its description.

The numerous, persistent floral bracts distinguish subsp. wagneri from
subsp. opegrapha and subsp. paquitensis. In late fruit, the bracts sometimes
fall, but even then, this subspecies can easily be recognized by the crowded
pedicel scars.

2c. Ardisia opegrapha Oerst. subsp. paquitensis (Lundell) Pipoly & Ricketson,
comb et stat. nov. (Figs. 15): Ardisia paquitensis Lundell, Phytologia 2:4.
1941. Graphardisia paquitensis (Lundell) Lundell, Phytologia 48:140. 1981. TYPE:
COSTA RICA. SAN JOSE: San José, low hills above Rfo Paquita, 5-50 m, 15 Aug.
1936 (fl, fr), CW Dodge & VE Goerger 9885 (HOLOTYPE: F, (F Neg. # 68245, LL-
TEX Neg. # 1971-81); ISOTYPE: MO, (LL-TEX Neg. # 1971-81).
Shrubs or small trees (0.6-)1-6 m tall. Branchlets 4—6(—7) mm in diam.
Leaves with blades elliptic to broadly elliptic 20.5-30 cm long, (7.5—-)8-

Sipa 18(2

es

Fic. 5. Ardisia opegrapha Oersted subsp. pa

flower, showing corolla and stamens. E.

quitensis (Lundell) Pipoly & Ricketson. A. Habit,
flowering branch. B. Detail of abaxial leaf surface. C. Detail of inflorescence. D. Detail of
Detail of stamen, showing adaxial and abaxial

surfaces. F. Fruit. A, C-E drawn from holotype. B, F drawn from isotype.

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 459

14.5 cm wide; petioles marginate, (1.5—)2—3(-3.5) cm long. Inflorescence
globose, a panicle of compound corymbs, (2.5—)3—5 (-7) cm long; peduncle
obsolete to 0.5(—1) cm long; inflorescence branch bracts 7.2-15.5 mm long,
2.6-6.5 mm wide, caducous; floral bracts 3.7—4.8 mm long, 0.8-1.3 mm
wide, caducous; pedicels 8.5-13 mm long. Flowers 7.5-10 mm long; se-
pals oblong, 4.5—-7 mm long, 2.5—3 mm wide; corolla 7—9.5 mm long, the
tube 0.6—0.7 mm long, the lobes widely ovate to oblong, 6.5—9 mm long,
4.6-5.7 mm wide; stamens 4—5,2 mm long, the staminal tube 0.6-0.8 mm
long, the filaments 1—2.5 mm long, 0.5—-0.7 mm wide, the anthers 3.3—
3.6 mm long, 0.9-1.1 mm wide at base; ovary with style 4-4.2 mm long,
the ovules 11-12, pluriseriate. Fruzt 4.5-5(—7) mm in diam.

Distribution —Endemic to Costa Rica, in the Provinces of Limén, Cartago,
San José and Puntarenas (especially common in the Cantons of Osa and
Golfito); sea level—1,400 m elevation.

Ecology and conservation status.—Subspecies paquitensis occurs in primary
and secondary tropical wet forests as a locally infrequent component of the
understory. Because of its restricted distribution, it should be considered
threatened.

Etymology.—The specific epithet ‘paquitensis’ refers to the type locality,
along the Rio Paquita, Province of San José, Costa Rica.

Additional specimens examined. COSTA RICA. Cartago: above Tuiz, road to Morravia
Chirripd, without elev., 1 Oct 1982 (fr), D. Hazlett 5101 (F, LL-TEX). Limén: Reserva
Indigena Talamanca Sukut, mouth of Rio Sukut at Rio Urén path to SE, toward Purisquf,
09° 24' 15" N, 82° 58' 10" W, 350-550 m, 6 Jul 1989 (fl), B. Hammel et al. 17551 (BRIT,
MO). Puntarenas: Canton Osa, Reserva Forestal Golfo Dulce Aguabuena, 08° 41' 50" N,
83° 30' 43" W, 40-50 m, 2 Aug 1991 (fl), R. Aguilar 178 (MO): Cantoén Golfito, Parque
Nacional Corcovado, Peninsula de Osa, Estacién Sirena, Sendero Ollas, 08° 28' 50'
83° 35' 30" W, 10 m, 13 Jun 1994 (fl), R es 3411 (BRIT, MO); Cantén Osa, vic
ity of Palmar Norte, along Rfo Grande de ‘Terraba, sea level, 2 Jul 1949 (fl), P Allen 5317
(A, F 3-sheets, GH, MO); vicinity Palmar Norte de Osa, 30 m, 6 Sep 1950 (fl), P Allen
5657 (F); Parque Nacional Corcovado, Peninsula de Osa, Sendero Mirador, 08° 33' 10"
N, 83° 30' 40" W, 200 m, 30 Aug 1995 (fr), L. Angulo 487 (MO); Reserva Bioldégia Carara
Estaci6n Quebrada Bonita, 09° 46' N, 84° 36' W, 30 m, 26 Jun 1990 (fl), &. Bello C. & E.
Rojas 2283 (BRIT, MO); near airport, 4 mi W of Rincén de Osa, 08° 42' N, 83° 31' W,
30 m, 4-7 Jun 1968 (fl), W. Burger & R. Stolze 5463 (F); Cantén Golfito, along hwy from
Rio Claro (on Panamerican Hwy.) to Golfito, 2.5 m SE of Golfito, 27.5 mi S of Rio Claro,
08° 36' N, 83° 04' W, 60 m, 15 Sep 1987 (fr), 7 Croat 67621 (BRIT, LL-TEX, MO);
Sirena, 08° 28' 00" N, 83° 35' 00" W, 1-200 m, 26 Jun 1991 (fl), P Delprete 5117 (TEX);
Quebrada Benjamin, Palmar Norte, 08° 59' N, 83° 28' W, 50 m, 14 Jan 1993 (fr), A
Gentry et al. 78808 (BRIT, MO); Quebrada Cunabri, Hitoy-Cerera, Baja Talamanca, 500
800 m, Jul 1984 (fl), L. Gémez 24093 (MO); divide between Quebrada Benjamin and
Quebrada Batambal, SW slope of Fila Retinto, NW of Palmar Norte, 08° 59' 00" N, 83°
28' 30" W, 300-400 m, 9 Jul 1990 (fl), 4. Grayum & R. Evans 9867 (MO); La Palma,
Rio Rincén headwaters, trail to Cerro de Oro, 08° 35' N, 83° 30' 40" W, 100 m, 30 Jul
1990 (fl), G. Herrera 4070 (BRIT, MO); Playa Cacao, lower basin of Quebrada Nazareno,
08° 37' 50" N, 83° 11' 00" W, 70 m, 27 May 1994 (fl), G. Herrera & G. Rivera 7081 (F);

460 Sipa 18(2)

between Palmar Norte and Puerto Cortéz, Osa Peninsula, 50 m, 6 Aug 1964 (fl), A. Kies

[ 39 (F); In Webb’s forest, between Palmar Sur and Puerto Cortéz, 400 m, 23 Aug 1‘
), A. Jiménez 3462 (F); Estacién Sirena, Naranjo Trail, 08° 27-30' N, 83° 33-38' o sea

oe 150 m, 27 May 1988 (fl), C. Kernan 537 (MO); Monkey Woods, 08° 27- 30' N,
83° 33-38' W, sea level, 16 Aug 1988 (fr), C. Kernan er P Phillips 828 (MO); Estacién
Sirena, S of Rio Sirena along Rio C amaronal, 08° 28' N, 83° 35' W, sea level, 22 Nov

1981 (fr), 8. Knapp 2177 (MO); Sue National Park, on ills 0-1 km W of the park
headquarters at Sirena, 08° 29' N, 83° 36' W, sea level-200 m, 4 Jul 1977 (fl), R. Liesner

2832 (MO); Along tri i" and in a bh ceween park hee idqu arters to Sirena and Pavo, 08°
30' N, 83° 36' W, sea level—10 m, 5 Jul 1977 (fl), &. Liesner & G. Vega 2900 (MQ); Puntarenas-
San José border, Baru, 100 m, 16 Aug | 974 (fl), P Maas & J. aie 1359 (F); Osa Pen-
insula, Coronado de Osa, 08° 32' 30" N, 83° 18' 30" W, 1-10 m, nie 1995 (fl), w
Madrigal 76 (MO); Cantén Osa, Fila Estero Guerra, Sierpe, 34' 30" N, 83° 34' 50"
W, 300 m, 27 Sep 1991 (fr), f, Marin 204 (MO); near the airport area, a mi W of Rincén
de Osa, 100 ft [30 m], 8 Aug 1967 (fr), 2 Raven 21627 (K, MQ); along road to Pacific
Ocean, W of Rincén de Osa, Osa Peninsula, near Mile 15, 100 ft [30 m}, ‘8 Aug 1967 (fl),
P Raven 21681 (F, MO); Rinceén de Osa, region N of airfield and ridges running NE,
100-300 m, 23 Jul 1974 (fl, fr), /. Usley & K. Utley 1194 (F); San Luis ae Turrubales, 450
m, 20 Jul 1933 (fl), MZ. Valerio 651 (I ). Sa n José: Cantén Pérez Zeledén, along road be-
tween San Isidro General and Dominceal, Fila Tinamastes, 09° 18' 24" N, 83° 46) 11" W,
990—1,100 m, 9 a 996 (fl), : Croat & D. Hannon 79101 (MO); Cantén Tarrazu, San
Lorenzo, 09° 34' 20" N, 84° 03' 52" W, 1,400 m, 9 Jul 1997 (fr), L. Gutiérrez et al. 64
(MO); fences : eee iG Vibora de Puriscal, 1,030 m, 23 Apr 1963 3 (fl), A. a Nez
692 (F); El General Basin, 675—900 m, Jul 1945 (fl), A. Skutch 5201 (F, MO, NY

“——

Ardisia opegrapha subsp. paquitensis is distinctive because of its oe
inflorescence, dwarfed by relatively large elliptic to broadly elliptic leaves.

3. Ardisia bartlettii Lundell

Shrubs or small trees (0.5—)2—10 m tall, 1—5 cm in diam. Branchlets slen-
der, (1.5—)2—3(-3.5) mm in diam., glabrous throughout. Leaves with blades
membranaceous to chartaceous, lanceolate elliptic or oblong 5.5—-16 cm
long, 1.8-7 cm wide, apically acute to acuminate, basally acute to acumi-
nate, decurrent on the petiole, midrib impressed above, prominently raised
below, the secondary veins prominulous above, inconspicuous below, densely
and prominently punctate and punctate-lineate, glabrous, the margins entire,
undulate to irregularly crenulate; petioles marginate to canaliculate, slen-
der to stout, 0.2—1.1 cm long, glabrous. /nflorescence terminal, bipinnately
paniculate, 2—6 cm long, 3.5—7 cm wide, the rachis glabrous, the branches
terminating in corymbs; peduncle obsolete to 0.5 cm; secondary inflores-
cence bracts early caducous, membranaceous, oblong, (2.8—)5.5—11.2(-14.2)
mm long, (1.3—)2.5—3.9(—5.4) mm wide, apically acute to obtuse, the midrib
prominulous above and below, the secondary veins inconspicuous, densely
and prominently black punctate, glabrous, the margins entire, sparsely elandular-
ciliolate; floral bracts very early caducous, unknown, at times not leaving
detectable scars axillant to pedicel (aborted at primordial stage ?); pedicels
6-15 mm long, slender, mostly accrescent, glandular-grandulose. Flowers

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 46]

5-merous, chartaceous, 6.8—9.2 mm long; calyx with sepals free, broadly
ovate, 1.5—2.5 mm long, 0.9—2 mm wide, apically acute, prominately punctate
and punctate lineate, glandular-grandulose within, opaque except at mar-
gin, the margins erose to subentire, ciliolate; corolla rotate, 6.5-9.2 mm
long, the tube 0.5-1.4 mm long, the lobes ovate, elliptic or lanceolate, 6-
8 mm long, 1.6—5 mm wide, apically rounded, prominently punctate and
punctate-lineate, yellow glandular-grandulose at base between corolla lobe
and tube junction and above staminal tube within, glabrous otherwise; margins
entire, ciliolate; stamens 3.5-5.7 mm long, the staminal tube 0.5—1.5 mm
long, the filaments 0.5—1.5 mm long, 0.2—0.5 mm wide at base, punctate,
glabrous, the anthers linear-lanceoloid, 2.5—3.5 mm long, 0.6-0.9 mm wide
at base, apiculate, basally sagittate, dehiscent by subapical pores, the con-
nective darkened; ovary glabrous, the style 4.5—-5 mm long, slender, stylopodic
at base or not stylopodic (tapered), punctate, glabrous, the ovules 13-16,
pluriseriate. Hruit globose to oblong, 4—5.2 mm long, 4.2—5 mm wide, densely
and prominently punctate, glabrous.

Because of its yellow glandular-granules within the corolla tube, Ardisia
bartlettii is most closely related to A. opegrapha, but is easily separated from
it by the chartaceous ovate, sepals 1.5—2.5 mm long, with acute apices, opaque
except at margin, and the margins subentire to erose. The glabrous fila-
ments are also distinctive. Ardisia bartlettii is comprised of two subspecies,
separated by size of floral parts, tapering vs. stylopodic style base, and habitat.

3a. Ardisia bartlettii Lundell subsp. rete (Figs. 1E, 6). Ardisia bartletii
Lundell, Contr. Univ. Michigan Herb. 7:37. 1942. pees bartlertii (Lundell)
Lundell, Phytologia 48:140. 1981. TYPE: et CANAL ZONE: Barro Colorado
Island, Gattin Lake, along William Morton Wheeler Tr ae ee elev., 8-10 Aug
1940 (fl, fr), A.A. Bartlett & T. Lasser 16720 (HOLOTYPE: MICH; ISOTYPES: LL- .
TEX, MO (LL-TEX Neg. # 1971-20)).

>

Ardisia romeroi Cuatrec., Revista. Acad. Colomb. Ci. Exact. 8:319. 1951. SYN. NOV. Graphardisia
romerot (Cuatrec.) Lundell, Wrightia 7:46. 1982. Type: COLOMBIA. Botivar: Camino
de Monte Libano a San Pedro, without elev., 28 May 1949 (fl), R. Romero Castafieda

756 (HOLOTYPE: F (LL-TEX Neg. 1971-92)).

Ardisia lewisti Lundell, Wrightia 4: 146, 1970. SYN. NOV. Graphardisia lewisti (Lundell)
Lundell, Bese 48:140. 1981. Type: PANAMA. CoLon: Santa Rita Ridge, ca.
5.5-6 mi E of Transisthmian Highway, without elev., 9 Apr 1969 (fl), WH. Lewis,
DM. ie L.H. Durkee & R.K. Baker 5377 (HoLotyre: LL-TEX; isorypes: LL-TEX,
MO, MOCZ n.v., UC).

saa tutrana Lundell, aaa 6:91. 1979. SYN. NOV. Graphardisia tuirana (Lundell)

indell, Wrightia 7:46. . Type: PANAMA, DariéNn: Rio Tuira and Rio Paca,
oe elev., 21 Jun ee A. Duke 5025 (HoLtotrype: LL-TEX; isorypes: BRIT,
D)

GH,

eee Lundell, oe 48:135. 1981. SYN. Nov. Graphardisia sapoana (Lundell)
Winer 7:46. Type: PANAMA. Darien: Cerro Sapo, NE slope of
summit, approach from ee knife edge ridge before ascent to top, 2,800 ft
[853 m], 9 May 1979 (fl), B. Hammel 7297 (HoLotyrE: LL-TEX; isoryPe: MO).

462 Sipa 18(2)

Fic. 6. Ardisia bartlettii Lundell subsp. bartlettii. A. Habit, flowering branch. B. Detail of
abaxial leaf surface. C. Detail of inflorescence. D. Detail of flower, showing corolla and

stamens. E. Detail of stamen, showing abaxial surface. E Detail of stamen, showing laterial
surface. G. Detail of stamen, showing adaxial surface. H. Fruit. >, E-H drawn from
holotype. D drawn from 7? Croat 15005.

PIPOLy AND RICKETSON, Revision of Ardisia subgenus Graphardisia 463

Shrubs (0.5—)2—10 m tall, 1—5 cm in diam. Branchlets slender, 1.5—3 mm
in diam., glabrous throughout. Leaves with blades 5.5-12.5 cm long, 1.8—
4.5 cm wide; petioles slender, canaliculate and slightly marginate, 0.2-1.1
cm long. /nflorescence 2-6 cm long, 4-6.5 cm wide; peduncle obsolete to
0.5 cm; secondary inflorescence bracts 5.5—-14.2 mm long, 2.5-5.4 mm
wide; pedicels 6-15 mm long, slender, accrescent, glandular-grandulose.
Flowers 6.8—7.2 mm long; sepals 1.5—1.8 mm long, 0.9-1.1 mm wide; corolla
6.5—7.7 mm long, the tube 0.5—1.2 mm long, the lobes ovate or elliptic to
lanceolate, 6—6.5 mm long, 1.6—4.1 mm wide; stamens 3.5—4.8 mm long;
the staminal tube 0.5—1.3 mm long, the filaments 1-1.5 mm long, 0.4—
0.5 mm wide at base, punctate, glabrous, the anthers 2.5—2.8 mm long,
0.6—0.8 mm wide at base; ovary glabrous, the style 4.5—4.8 mm long, slen-
der, not stylopodic at base, punctate, glabrous, the ovules 15, pluriseriate.
Fruit globose to oblong, 4.5—5.2 mm long, 4.2—4.5 mm wide, conspicu-
ously punctate, glabrous.

Distribution —Known only from eastern Panama (Coclé, Colén, Panama,
Darién, San Blas), and on the Caribbean coast of Colombia (Bolivar), growing
at sea level—900 m elevation.

Ecology and conservation status —Ardisia bartlettii subsp. bartlettii occurs
along streams and rivers in wet and pluvial lowland, premontane and cloud
forests. It appears to be relatively tolerant of disturbance, so we believe it is
not threatened at this time.

Etymology.— The specific and subspecific epithet honors Harley Harris
Bartlett (1886-1960), American botanist at the University of Michigan at
Ann Arbor, who collected the type specimen.

Additonal specimens examined. PANAMA. Coclé: Santa Rita Ridge, 5-10 mi from
Transisthmian Hwy., sea level-100 m, 28 Jun 1984 (fl), ane bie 15548 (BRIT, LL-
TEX, MO, PMA); from Torti to Pilota del Toro, mountain overlooking Torti Arriba, without
elev., 27 Aug 1977 (fr), J. Folsom e G. Alonzo de Monte 4973 (LL- ate MO), 5012 (MO);
mountains above Torti Arriba, without elev., 2 Dec 1977 (fr), /. Folsom et al. 6576 (MO);
4—5 hours walk upriver from Torti Arriba, 200-300 m, 6 Dec 1977 (fr), /. Folsom et a
6844 (MO); lowland area ane ae stream that passes through Torti, on the S side o
Pan-American Hwy., on Quip r river, without elev., 25 ee 1980 (fl), /. Fan
etal. 7825 (FE, TEX); Hill S ae uted 8 km E of Ipeti, 08° 55' N, 78° 20' W, 600 m, 18
Sep 1982 (fr), C. Hamilton & W. D'Arcy 1387 (LL-TEX, MO); Bipclinc road, N pe
upstream of the tenth bridge (Rio Guacharo), beyond the big waterfall, 09° 10' N, 79° 45'
W, 100 m, 4 Aug 1984 (fl), G. de Nevers & D,. Hews 3627 (LL-TEX, MO); Pipeline Road
near Gamboa, Rio Mendoza, upstream of road, without elev., 09° 09' N, 79° 42' W, 7 Jul
1985 (fl), G. de Nevers 5963 (LL-TEX, MO, US). Colén: Trail from end of Santa Rita
Ridge Road to Rio Piedras, 1,400—1,600 ft [427-488 m], 23 Mar 1980 (fr), 7) Antonio
3889 (LL-TEX, MQ); Santa Rita Ridge, along road, ca. | mi from Boyd Roosevelt Hwy.,
without elev., 9 Jul 1971 (fl), 7’ Croat 15340 (MO); Santa Rita Lumber Road, ca. 15 km
E of Coldn, without elev., 20 Apr 1971 (fl), R. Dressler & N. Williams 3967 (MO); Santa
Rita Ridge, 20.7 km from Transisthmian Hwy., 09° 23' N, 79° 40' W, 530 m, 22 Mar
1992 (ster.), R. Foster et al. 14038 (MO); ridge top 1-3 mi W of Portobello, without elev.,

—

464 Sipa 18(2)

7 Sep 1971 (fr), A. Gentry 1758 (F, MO); Santa Rita Ridge, ca. 12 km from Transisthmian
Hwy., without elev., 28 Jun 1978 (fr), B. Hammel 3646 (MQ); end of Santa Rita Ridge
Road, 21 km from Transisthmian Hwy., 09° 25' N, 79° 37' W, 400-500 m, 22 May 1982
(fl), S. Knapp & R. Schmalzel 5247 (LL-TEX, MO); 26-28 km from Transisthmian Hwy.
on the Santa Rita Ridge Road, 09° 25' N, 79° 37' W, 250-400 m, 23 May 1982 (fl), S.
Knapp e R. Schmalzel 5268 (LL-TEX 2-sheets, MO); Santa Rita es Road, 21—26 km
from Transisthmian Hwy., 09° 25' N, 79° 37' W, 500-550 m, 4 Jul 1982 (fl, fr), S. Knapp
5892 (LL-TEX, MO); Santa Rita Ridge, SE of Coldn, along rodge road, 10-12 mi from
Transisthmian Hwy., 09° 20' N, 79° 45' W, 550 m, 21 May 1986 (fl), G. McPherson 9206
(MO); Santa Rita Ridge Road, ca. 6 km from Boyd Roosevelt Hwy., without elev., 26 Sep
1974 (fr), S. Mori & J. Kallunki 2140 (LL-TEX, MOQ); Santa Rita Ridge road, 20-25 mi
from Transisthmian Hwy., 09° 24' N, 79° 39' W, 100-1,200 m, 10 Oct 1980 (fr), K. Sytsma
1523 (LL-TEX, MO); Upper Rio Piedras headwaters, along trail from end of Santa Rita
Ridge Road, ca. 11 km SW of Cerro Braja, 09° 25' N, 79° 35' W, 600-700 m, 30 Apr
1981 (fl), K. Sytsma et al, 4192 (LL-TEX, MOQ); along Santa Rita Ridge, road to Estacién
Calibar Lluvia el Agua Clara, 09° 22' N, 79° 42-45' W, 1,300 ft [396 m], 26 Jun 1971
(fl), G. Webster & R. Dressler 16745 (MO). Darién: vicinity of Torti, 38.6 mi E of Bayano
Dam Bridge, trail from village to mountains near Rio Torti, 50-250 ft [15-76 m], 17
May 1980 (fl), 7° Antonio 4674 (LL-TEX, MQ); Rio Tuira 2 mi upstream from Boca del
Cupa, 4 Aug 1962 (fr), /, Duke 5381 (MO); vicinity of Las Pifias, without elev., 2 Mar
1967 (fr), /. Duke 10612 (LL-TEX); S of Garachiné near Pacific coast above Casa Vieja,
along boundary trail of Parque Nacional Darién, W flank, Serranta del Sapo, 07° 58' N,
78° 23' W, 50-150 m, 21 May 1991 (fl), N. Hensold 1078 (K MQ); Casa Vieja hacia
Cerro Sapo, 07° 58' N, 78° 23’ W, 180-500 m, 23 May 1991 (fl), H. Herrera, et al. 984
(BRIT, KF MO, PMA); S of Garachiné on W flank of Serrania del Sapo, above place called
Casa Vieja, along boundary trail of Darién National Park, 07° 58' N, 78° 23' W, 300-500
m, 24 May 1991 (fl, fr), G. McPherson 15369 (BRIT, MO). Panama: Barro Colorado,
Mojinga swamp near mouth of Rio Chagres, 1 m, 11 Mar 1935 (fl), P Allen 865 (MQ);
road to Cerro Jefe, 900 m, 9 Jul 1976 (fl), T. Antonio 1296 (BRIT, LL-TEX, MO); Barro
Colorado Island, without elev., 1931 (fl), S. Aviles 10 (MO); Barro Colorado, Snyder-Molina
Trail, without elev., 29 Jun 1940 (fl), 7. aha 4796 (F); near Gamboa, without elev.,
26 Jun 1972 (fl); A. Clewell & E. Tyson 3222 (MQ); Barro Colorado, Drayton Trail, with-
out elev., 22 May 1968 (fl), 7) Croat 5756 (BRIT, MO); Barro Colorado, Wheeler Trail,
without elev., 19 Sep 1968 (fr), 7) Croat 6215 (BRIT 2-sheets, MO); Barro Colorado,
Wheeler Trail, without elev., 11 Jun 1970 (fl, fr), 77 Croat 10845 (F, MO); Barro Colo-
rado, Shannon Trail, without elev., 15 Jul 1970 (fl, fr), 7 Croat 11271 (MO); Barro Colo-
rado, Wheeler Trail, without elev., 17 Sep 1970 (f], fr), 7? Creat 12219 (MO); Barro Colo-
rado, without elev., 20 May 1971 (fl), 7) Creat 14650 (MQ); Barro Colorado, Lake Trail,
without elev., 16 Jun 1971 (fl), 7 Croat 15005 (MO); Barro Colorado, Barbour Trail,
without elev., 25 Jun 1971 (fl), 7 ee 15105 (K MQ), between Fort Sherman and Fort
San Lorenzo, without elev., 10 Jul 1971 (fl), 7 Croat 15414 (LL-TEX, MO); Cerro Jefe,
2,900 ft [884 m], 21 Jul 1972 (fl), 7. a eA. Gentry 10249 (F, MQ); Barro Colorado,
Snyder Molino Trail, along edge of trail, 300-600 m, 25 Jun 1960 (fl), J. Ebinger 182
(BRIT, MO); along the border of Canal Zona-Colén Prov., from parallel in NW shore of
Canal along road to Achiote, within 2 mi of Achiote, without elev., 23 Jun 1977 (fl), /-
Folsom 3880 (LL-TEX, MO); road leading around the shoulder of Cerro Jefe, turns to the
right at 21.5 km from the Pan-American Hwy., around the first stream encountered, 800
m, 29 Sep 1977 (fl, fr), J. Folsom et al. 5663 (MO); ca. 1 mi ss from Frizzel’s Finca
Indio, om slope of Cerro Jefe, without elev., 9 Sep 1970 (fl), R. Foster & H. Kennedy 1849
(LL-TEX); Barro Colorado Island, without elev., 9 Oct 1985 (fr), N Garwood e §. Arne
1595 (F), 1 Dec 1985 (fr), N. Garwood 1687 (F), 28 Sep 1986 (fl, fr), N. Garwood 1943

“~~

rc

WS

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 465

(F), 18 Nov 1988 (fr), N. Garwood 2658 (F); Pipeline Road, without elev., 30 Sep 1971
(fl), A. Gentry & R. Dressler 1979 (MO), 6 mi N of Gamboa, without elev., 28 Dec 1971
(ster.), A. Gentry 3319 (MO); Cerro Jefe, region roadside along road to Altode Pacora, 2
km N of turnoff to Cerro Jefe radio tower, along small stream, 2,500-2,700 ft [762-823

m], 30 Sep 1978 (fl), B. Hammel 4854 (MQ); Fort Sherman, Atlantic coastal forest, _
out elev., Jul 1965 (fl), 4. Hayden 95 (MQ); trail along Rio Petitpie from road to Fort
Shennan from Gatum Locks, without elev., 22 Oct 1974 (fr), 8. Mori & J. Kallunki ba
(MO), 12 Dec 1974 (fr), S. Mori & J. Kallunki 3661 (MO); ca. 12 km NW of Gamboa,
without elev., 26 Aug 1975 (fr), S. Mori 7943 (LL-TEX, MO), 9 km NW of Gamboa,
150 m, 29 Oct 1973 (fr), M. Nee 7657 (LL-TEX, MO), 125 m, 28 Nov 1973 (fl), AZ. Nee
8435 (LL-TEX 2-sheets, MO); Along road $1, 4 km W of Gatun Dam, 190-200 m, 20
Dec 1973 (fl), M4. Nee 8913 (MO 2-sheets); Barro Colorado, near A.V. Armour Trail, 5,
on Hubbell Permanent Plot, without elev., 13 Nov 1981 (fl), R. Schmalzel & M. Aide 56
(MO), Sugar Mill, without elev., 20 Dec 1931 (fr), O. Shattuck 611 (F, MO), Standley
No. 20, without elev., 31 Jul 1934 (fr), O. Shattuck 1098 (BRIT, EK MO); Gatun Lake,
120 m, 18-24 Nov 1925 (fr), P Standley 41029 (LL-TEX), Barro Colorado, Zetele Trail,
without elev., 6 Jul 1931 (fl), D. Starry 28 (F, MO); Pefia Blanca Trail, without elev., Jul
1931 (fl), D. Starry 178 (F); Cerro Jefe, 850-900 m, 29 Oct 1980 (fr), K. Sytsma 1975
(BRIT, MO); Cerro Jefe, 1.5 mi down right turnoff, 6.7 mi past Goofy Lake
Dec 1980 (fl, fr), A. Sytsma et al. 2900 (MQ); Cerro Jefe, 6 mi past Cerro Azul on road to
Altos Pacora, 2,600 ft [792 m], 19 Feb 1981 (fr), K. Sytsma e& W. D’Arcy 3690 (LL-TEX,
MO); vicinity of Salamanca Hydrographic Station, Rio Pequenf, 80 m, 28-29 Jul 1938
(fl), R. Woodson et al. 1569 (F, MO). San Blas: Nusagandi, Campo de PEMASKY, ca. 20
km on EI Llano-Carti Road, trails near station, 09° 18' N, 78° 59' W, 350 m, 1 May 1992
(fl, fr), R. Foster et al. 917 (MO); 3-4 hours up Rio Mulatupo by foot, without elev., 17
Aug 19067 (fr), J. Kirkbride 216 (MO); Udirbi Reserve, waterfall trail along park bound-
ary, 09° 18' N, 78° 58' W, 350-400 m, 21 Jul 1986 (fl), /. ee et al, 244 (MO); El
Llano-Carti Road, 24.5-25 km from Inter-American Hwy., near continental divide, 13
Apr 1975 (fl), S. Mori & J. Kallunki 5556 (MO), 17.4 km . Inter-American Hwy, 09°
19’ N, 78° 55' W, 350 m, 27 Sep 1984 (fl, fr), G. de Nevers 3943 (BRIT, MO), km 16.7,
trail W to waterfall 5 km from road, 09° 19' N, 78° 55' W, 350 m, 16 Jun 1985 (fl), G. de
Nevers . S. Charnley 5899 (LL-TEX, MO, US); Cerro Habu, trail from Rio Sidro, 09°

8° 49' W, 800-1,400 ft [244-427 m], 18 Dec 1980 (fr), A. Sytsma 2650 (MO).

SOnteae.. misinterpreted Ardisia romerol aSa member of subgenus lcacorea,
largely because of the early caducous floral bracts and concolorous, poricidally
dehiscent anthers. However, it closely matches the type of Ardisia bartlettii
subsp. bartlettii. Ardisia lewisii is notable only for its inflorescence with pedicels
in loose (low) anthotactic spirals. Ardisia tuirana was separated because of
its somewhat less punctate, ovate sepals and sparse, finely stellate tomen-
tum on the lower branches of the inflorescence. It is otherwise not signifi-
cantly different from the type of the subspecies. Populations correspond-
ing to the type of Ardisia sapoana vary from the subspecies type only by
having sessile inflorescences with longer secondary inflorescence branch bracts,
and longer corolla tubes.

Ardisia bartlettii subsp. bartlettii may be separated from subspecies /ilacina
by the shorter calyx and corolla, the generally shorter stamens and tapering
style base.

466 Stpa 18(2)

3b. Ardisia bartletii Lundell subsp. lilacina (Lundell) Pipoly & Ricketson,
comb. et stat. nov. (Figs. 1F, 7). Ardisia lilacina Lundell, Wrightia 3:198. 1966.
Graphardisia lilacina (Lundell) Lundell, Phytologia 48:140. 1981, TYPE: PANAMA.
OLON: Porto Belo [Bello], beach, without elevation [ca. sea level], 13 Jul 1964 (fl),

7, Dwyer 4354 (HOLOTYPE: MO

Shrubs or small trees 0.5-)2-7 m tall, 1-4 cm in diam. Branchlets slen-
der, 2—3.5 mm in diam., glabrous throughout. Leaves with blades 6—16 cm
long, 3.5—7 cm wide; petioles stout, marginate, 0.4—1 cm. /nflorescence 2.5—
4 cm long, 3.5—-7 cm wide, sessile or nearly so; peduncle obsolete to 0.2
cm; secondary inflorescence bracts 2.8-11.2 mm long, 1.3—3.9 mm wide;
pedicles 10-15 mm long, slender, glandular-glandulose. Flowers 8—9.2 mm
long; sepals 2—2.5 mm long, 1.5-2 mm wide; corolla 7.5-9.2 mm long,
the tube 0.5—1.4 mm long, the lobes ovate to elliptic, 7-8 mm long, 4.5—
5 mm wide; stamens 4.5—-5.7 mm, the staminal tube 0.5—1.5 mm long,
the filaments 0.5—1 mm long, 0.2—0.3 mm wide at base, punctate, glabrous;
the anthers 3—3.5 mm long, 0.7—0.9 mm wide at base; ovary glabrous, the
style 4.5-5 mm long, slender, stylopodic at base, punctate, glabrous, the
ovules 14-16, pluriseriate. Fruzt globose, 4-5 mm in diameter, conspicu-
ously punctate, glabrous.

Distribution.—Subspecies /ilacina is restricted to the Caribbean coast of
Panama (Colén, Panama, San Blas), with one disjunct population in the
Chocé, Colombia. We may also expect subsp. /lacina in lowland Antioquia
and Valle Departments of Colombia, areas sorely lacking in collections. It
grows from sea level-180 m elevation.

Ecology and conservation status.—This subspecies occurs in strand veg-
etation and beach forests. It is said to be locally uncommon, so it should be
considered threatened.

Etymology.—The epithet ‘lilacina’ refers to the striking lilac color of the
flowers.

Additional specimens examined. PANAMA. Coldén: vicinity of Viento Frio, along the
beach, sea level, 07-08 Aug 1911 (fl), 1. Pittier 4114 (F); Rio Indio de Faté, sea level, 24
Aug 1911 (fl), H. Prttier 4273 (F, LL-TEX, NY); 3 km SW of Rio Guanche along road
from Puerto Pilén to Portobelo, sea level, 20 May 1981 (fl), « Syisma er L, —
4791 (LL-TEX, MO). Panama: Rio Providencia and ridge S of river, 5-170 m, 5 Dec
1973 (fl, fr), A. Gentry & M. Nee 8706 (LL-TEX, MO). San Blas: Rio C aoa pueblo
Cangandi, path to Quebrada Inadi, 09° 27' N, 79° 07' W, 50 m, 20 May 1987 (fl), H.
Herrera & P Pérez 127 (MO); Meds Pacifica de la Cordillera de San Blas, Cabecera del
Rio Piriadf, 09° 11' 05" N, 78° 16' W, 100-150 m, 14 Jun 1994 (fl), H. Herrera 1633
(BRIT, kK MO, PMA); Comarca de San Blas, tierra firme de Playén Chico, vicinity of the
aqueduct trail, 09° 17' N, 78° 15' W, 30 m, 12 Sep 1994 (fr), H. Herrera 1839 (BRIT,
MO, PMA); Playén Chico and vicinity San Blas, Yantuppu, coral island, without elev., 8
Jul 1975 (fl), & Stier 193 (MO). COLOMBIA. Chocé: Mpio. Acandi, Vereda El paramo,
Quebrada Sardi, 08° 20' N, 77° 06' W, 180 m, 22 May 1989 (fl), k. Fonnegra et al. 2792
(HUA, MO), - Rolddn et al. 1190 (HUA, MO).

PIPOLY AND RICKETSON, Revision of Ardisia subgenus Graphardisia 467

Fic. 7. Ardisia bartlettii Lundell subsp. /i/acina (Lundell) Pipoly & Ricketson. A. Habit,
naa branch. B. Detail of abaxial leaf surface. C. Detail of inflorescence. D. Detail of

wer. E. Detail of stamen, showing abaxial surface. F. Detail of stamen, showing laterial
en G. Detail of stamen, eee aa adaxial surface. H. Fruit. A-G drawn from holo-
type. H drawn from & Stier

468 Sipa 18(2)

Ardisia bartlettii subsp. lilacina may easily be separated from subsp. bartletii
by its longer perianth parts, usually longer stamens and unique stylopodic
(swollen) style base. While the stylopodic style base is unique within the
subgenus, there is no corresponding unique feature for subspecies bartlettii
to define it. Therefore, the two taxa are recognized at the subspecific level.
In addition, it is restricted to strand and beach forests, while subsp. bartlettii
is from tall wet forests.

EXCLUDED NAMES

Ardisia albovirens Mez, Repert. Spec. Nov. Regni Veg. 16:311. 1920. Graphardisia
albovirens (Mez) Lundell, Wrightia 7:46. 1982. TYPE: BRAZIL. ACRE: Rio Acre bei
Xapury [Xapuri], without elev., Jan. 1911 (fl), £. Ule 9682 (HOLOTYPE: B-destroyed
in 1943 (P Neg. # 4876); LECTOTYPE, here designated; K; photo of lectotype: LL-
TEX Neg. # 71-157).

Ardisia mgpniren), EF. Macbr., Candollea 5:397. 1934. SYN. Nov. Graphardisia nigrovirens
J. E Macbr.) Lundell, Wrightia 7:46. 1982. Type: PERU. Loreto: Puerto sins
Yurimaguas, lower Rio Huallage, 155-210 m, 15 Nov 1929 (fr), L. Williams 5081
(HOLOTYPE: F (LL-TEX Neg. # 1971-74)).

B)

Now that recent collections have scale us with material both in flower
and fruit, it is clear that this species is not a member of subgenus Graphardisia,
but rather, subgenus Ardisia. This species has ovate anthers with subapical
pores opening into slits, has panicles with racemose branchlets, inflores-
cence and secondary inflorescence branch bracts that are subcoriaceous to
coriaceous and cucullate, and coriaceous sepals that are quincuncial. We
postulate that the rather large floral and secondary inflorescence branch
bracts of the species may have led to its misplacement.

ACKNOWLDEGMENTS

We thank the Missouri Botanical Garden and the Flora Mesoamericana
Project, for funding that allowed J. Ricketson (MO) to travel to Fort Worth,
where he was joined by J. Pipoly (BRIT) on visits to the C.L. Lundell Herbarium
(LL-TEX), eee at the University of Texas at Austin. Without access to
that critical collection, assembled by C.L. Lundell over a period of nearly
60 years, the present study would not have been possible. We also thank
the curators of the herbaria cited for loans of specimens. We are grateful to
the staff of TEX, especially Billie L. Turner, Tom Wendt, Carol Todzia, Beryl!
Simpson, and José Panero, for their cooperation and hospitality. We are
also grateful to those who have been so instrumental in assisting us in our
work, including K. Gandhi (GH), Dan Nicolson (US), Gerrit and Jeany
Davidse, Linda Oestry, Mary Bard, and Catherine Mayo, (MO), Barney
Lipscomb, Lindsay Woodruff, Marjorie Weir and Jim Rivers (BRIT). We
are deeply grateful to Yvette “Tivvy” Harvey (K), G. Wagenitz (GOET),
Laurent Gautier (G), Asuncién Cano (USM), Carlos Reynel (MOL), Fred

PIPOLy AND RICKETSON, Revision of Ardisia subgenus Graphardisia 469

Barrie (F), Krzysztof Swierkosz (W/RSL), Franz Schuhwerk (M), Bruno Walnéfer
(W), and Karen Stilwell (BM) for searching for duplicates of type material.
Illustrations were prepared by the junior author, and we thank Ms. Pauline
Lawson, General Manager of The Copy Stop of Fort Worth, Texas for her
generous technical assistance with preparation of the plates for scanning.
Reviews of the manuscript by Michael Grayum and Charlotte Taylor, and
meticulous copy editing by Barney Lipscomb, greatly improved the pre-
sentation of the paper.

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brief history of the subject. Clarendon Press, Oxford, U.K. Pp. 54-62.

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NUMERICAL LIST OF TAXA

1. A. weberbaueri Mez 3. A. bartlettii Lundell
2. A. opegrapha Oerst. 3a. subsp. bartlettii
2a. subsp. opegrapha 3b. subsp. /idacina (Lundell) Pipoly & Ricketson

2b. subsp. waeneri (Mez) Pipoly & Ricketson
2c. subsp. paquitensis (Lundell) Pipoly &

Ricketson

LIST OF EXSICCATAE

The figures in parentheses refer to the numbers from the numerical list
of taxa. Collection numbers in boldface type indicate ype specimens.
Acevedo, D. et al. 128 (2b). Aguilar, R. 178 (2c); 3411 (2c). Aguilar, S. & A. Castillo 2116 (2a).
Allen, P. 72 (2a); 786 (2a); 865 (3a); 2226 (2a); 2571 (2a); 2577 (2a); (2a); 3535 (2a); 3561
(2a); 4404 (2a); 5317 (2c); 5657 (2c). Allen, P. & A. Alston 1846 (2a). Angulo, L. ee ae 487
(2c). Antonio, I. 1296 (3a); 2573 (2a); aes 1); 4674 (3a); en 1 (2a). Araquistain, M.. 7 (2b);
3087 (2b); ae a); 3132 (2b). Araya, F 147 (2b). Araya, F. et al. 306 (2b). Atwood, J. SR
Aviles, S. 10
Barbosa, C. 6412 (2a). Barringer, K. & L. Diego ew 1652 (2a). Bartlett, H. & T. Lasser en
(3a). Beck, C. s.n. (2a). Beck, S. 16323 (1). Bello C. 2, 291 (2b); 1032 (2b); 2929 (2b). Bello C. I
& E. Rojas 2283 (2c). Blum, K. et al. 2382 (2a). ote A. 4313 (2a); 6349 (2a); 12653 (2b); 19942
(2a). Burger, W. &

).

—
—

De
a);

eS

—

aa

).t
R. Baker 9847 (2b). Burger, W. & R. Stolze 5463 (2c). Burger, W. et al. 11684

Carballo, G. 151 (2b); 436 (2b). Carvajal, A. 269 (2a). Chacén, J. & A. Chacon 2153 (2b). Chrysler,
96 (3a). Churchill, H. 5687 (2a). Churchill, J. 82022 (2a). Clewell, A. & E. Tyson 3222 (3a).
Correa, M. et al. 5139 (2a). Croat, T° ee (Ga); ares a); 10845 (3a); 11271 (3a); 12204 (3a);
12219 (3a); 14650 (3a); 14746 (2a); 15005 (3a); 105 (3a); 15340 (3a); 15414 (3a); 25189 (2a);
nea 27350 (2a); 34205 (2a); 36349 (2 >); pene 46928 (2b); 66873 (2a); 67621 (2c).
Cr oat, 1. & J. Folsom 33865 (2a); 34267 (2a). Croat, T. & D. Hannon 79101 (2c). Croat, T. & G.
Zhu 76926 (2a).

Pipoty AND RIcKETSON, Revision of Ardisia subgenus Graphardisia 471

D'Arcy, W. 9535 (2a); 10274 (2a); 15019 (2a). D’Arcy, W. & C. Hamilton 14976 (2a). D’Arcy, W.
et al. 15548 (3a). Davidse, G. 24159 (2a); 24176 (2a); 24428 (2a). ee P. 5117 (2c). Dodge,

& V. Goerger 9885 (2c). Donnell Smith, J., 6677 (2b). Dressler, R. & N. Williams 3967 (3a).
Diep 5025 (3a); 5381 (3a); 8650A (2a); 10612 (3a); 15799 (2a). Dwyer, J. 1612 (2a); 2363 (2a);
4354 (3b). Dwyer, J. & A. Gentry 10249 (3a). Dwyer, J. & J. Kirkbride 7818 (2a).

Ebinger, J., 182 (3a); 751 (2b). Echeverria, J., 4147 (2a). Endres, A., s.n. (2a).

Folsom, J. 3880 (3a); 3975 (2a); 4029 (2a); 4925 (2a). Folsom, J. & G. Alonzo de Monte 4973 (3a);
5012 (3a). Folsom, J. & L. Collins 1613 (2a). Folsom, J. et al. 3375 (2a); 5663 (3a); 6844 (3a); 6991
(2a); 7825 (3a). Fonnegra, R. et al. 2792 (3b). Foster, R., 1600 (2a). Foster, R. & H. Kennedy, 1849
(3a). Foster, R. et al., 917 (3a); 14038 (3a). Friedrichsthal, E., s.n. (2b); 1296 (2b).

Garcia, T. 203 (2b). Garwood, N. 1687 (3a); 1943 (3a); 2658 (3a). Garwood, N. & S. Arne 1595
(3a). Cock N. et al. 961 (2b). Gentry, A. 1758 (3a); 3319 (3a). Gentry, A. & R. Dressler 1979
(3a). Gentry, A. & J. Dwyer 5512 (2a). Gentry, A. & S. Estensoro 70712 (1). Gentry, A. & M. Nee
8706 (3b). Gentry, A. et on 43897 (2b): 78808 (2c). Gomez, L. 22966 (2b); 24093 (2c). Gémez, L.
etal., 21982 (2a). Gémez-Laurito, J., 10215 (2b). ea M., 2403 (2b). Grayum, M. & R. Evans
9867 (2c). Grayum, M. & B. Hammel 5638 (2a). Grayum, M. & P. Sleeper 6103 (2a). Grayum, M.
et al., 8325 (2a). Grijalva, A. & P. Moreno, nes (2b); ee (2b). Grijalva, A. & D. Rios, 3493 (2b).
Grijalva, A. et al., 3380B (2b). Gudifio, E. 355 (1). Gutiérrez, L. et al. 64 (2c).

Haber, W., 1427 (2b); 1428 (2b); 2000 (2b); 2079 (2b); 2187 (2b); 2188 (2b); 2189 (2b); 2190
(2b); 2191 (2b); 2192 (2b); 2193 (2b); 2194 (2b); 2195 (2b); 2196 (2b); 2197 (2b); 2198 (2b);
2199 (2b). 2976-(2b) 2315: 2b)s 2316 (2b); 2317 (2b)e 2318 Ob)-2319\ Ob). 2320 (2b): 2321
(2b); 2322 (2b); 2323 (2b); 2324 (2b); 2325 (2b); 2326 (2b); 2327 (2b); 2329 (2b); 2331 (2b);
2439 (2b); 2440 (2b); 2441 (2b); 2442 (2b). Haber, W. & E. Bello C. 1995 (2b); 2667 (2b); 2691
(2b); 2693 (2b); 2888 (2b); 5031 (2b); 5039 (2b); 5055 (2b); 5063 (2b); 5641 (2b); 7439 (2b);
7454 (2b). Haber, W. & J. Wolfe ao ee Haber, W. et al. 1770 (2b); 11260 (2b). Hamilton, C.
& W. D’Arcy 1387 (3a). Hamilton & K. Krager 3973 (2a). Hammel, B. 1764 (2a); 3646 (3a);
4854 (3a); 7297 (3a); 11619 (2b); | i (2b);13761 (2a). Hammel, B. & M. Grayum 14201 (2a).
Hammel, B. et al. 17551 (2c). Hartshorn, G. 1254 (2b). Hayden, M. 95 (3a). Hazlett, D., 5064
(2a); 5101 (2c); 5229 (2a). Hensold, N., 1078 (3a). Herrera, G. 1060 (2b); 1435 (2b); 4070 (2c).
Herrera, G. y G. Rivera 7081 (2c). Herrera, H. 1633 (3b); 1839 (3b). Herrera, H. & P. Pérez 127
(3b). Herrera, H. et al. 984 (3a). Holm, R. & H. Iltis 315 (2b)

I] INBIO 53 (2b). Isern, J. 2234 (1).

Jacobs, B. 2871 (2b). Jaramillo, J. & A. Grijalva 11402 (1). Jiménez, A., 692 (2c); 2239 (2c); 3462
(2c); 3966 (2a).

Kernan, C. 537 (2c). Kernan, C. & P. Phillips 828 (2c). Kirkbride, J. 216 (3a). Kirkbride, J. & J.
Duke 1642 (2a). Klug, G. 3978 (1). ee S. 2177 (2c); 5892 (3a). Knapp, S$. & R. Schmalzel
5247 (3a); 5268 (3a). Knapp, S. & K. Sytsma 2498 (2a); 2518 (2a). Kress, W. & T. a ae 4266 (2a
Kress, W. & T. Prinzie, 4572 (2a); 4582 ae Kupper, W., 1529 (2b

Laguna, A. 27 (2a). LeDoux, D. 2630 (2a). Lehmann, F. s.n. (2b). Lent, R. 1321 (2a); 1765 (2a);
2693 (2a). Lewis, W. et al. 1803 (2a); 3038 (2a); 5377 (3a). Liesner, R. 2832 (2c). Liesner, R. & G
Vega 2900 (2c). Lobo, M. 25 (2b). Luteyn, J. 4012 (2a)

Maas, P. & J. Cramer 1359 (2c). Madrigal, M.. 76 (2c). Marin, J. 204 (2c). McDaniel, S. 6912 (2a).
McDonagh, J. et al. 244 (3a). McDowell, T. 562 (2b); 1007 (2b). McPherson, G. 7458 (2a); 9206
(3a); 9248 (2a); 10744 (2a); 10933 (2a); 12797 (2a); 13513(2a); 15369 (3a). Méndez, R. 45 (2a);
47 (2A); 80 (2a). Mendoza, R. 26 (2a). Miller, J. & J. Sandino 1257 (2a). Molina, A. et al. 17708
(2b). Morales, J. 318 (2a). Moreno, P. 7730 (2b); ee (2b); 17273 vee 17312 (2b); 19119 (2b);

19225 (2a); 20597 (2b); 26297 (2a). Moreno, 2. & Robleto 20745 (2b); 25981 (2b). Moreno, P.
& J. Sandino 13039 (2b); 13100 (2b). Mori, S. Ba (2a); 7034 (2a); 7943 (3a
Mori, S. & J. Kallunki 2140 (3a); 2520 (2a); 2665 (3a); 3661 (3a); 5556 (3a). tou S. et al. 7545
(2a). Tp H., 1248 (2a).

Nee, M., 7657 (3a); 8435 (3a); 8913 (3a); 28425 (2b). Nee, M. et al. 35424 (1). Neill, D. 4478
(2b). Nevers, G. de 3943 (3a); 5963 (3a). Nevers, G. de & S. Charnley 5 hee (3a). Nevers, G. de &

aauity,

472 Sipa 18(2)
D. Hews 3627 (3a). Nufiez, P. et al. 8053 (1).
Oersted, A. 29 (2a); 29A (2a). Ortiz, F. 49 (2b); 112 (2b); 201 (2b); 281 (2b); 324 (2b); 2130 (2b).

Sa , J. 3516 (2b); 3670 (2b); 4420 (2b); 5010 (2b). Pipoly, J. etal. 12168 (1). Pitman, N. & M. Aulestia
2 (1). Pittier, H. 2298 (2a); 4114 (3b); 4273 (3b); 11190 (2a). Proctor, G. et al. 27151 (2a).

Raven, P. 21627 (2c); 21681 (2c); 21889 (2a). Rios, D. 232 (2b). Rios, P. 385 (2b). Rivera, G. 1452

(2b). aon F et al. 1190 (3b). Romero Castafteda, R. 1756 (3a). Ruiz, M. et al. 165 (2a); 210

(2a); 5139 (2a); 7320 (2a); 7800 (2a); 7865 (2a). Russell, G. et al. 870 (2b)

—

Salick, J. 7860 (2b). Sandino, J. 160 (2b); 186 (2b); 192 (2a); 3260 (2b); 3369 (2b); 3427 (2b);

4534 (2a). Schmalzel, R. & M. Aide 56 (3a). Schubert, B. 1086 (2b). Schunke V., J. 3983 (1); 4039

(1); 4983 (1); 6449 (1); 7647 (1); 8133 (1); 8384 (1). Seeman, B. C. s.n. (2b). Seibert, R. 456 (2a).

Seidel, R. & V. Baptista 2824 (1 (1). Seidel, R. & M. Schulte 2316 (1). Seidel, R. et al. 2671 (1). Seymour,

FE. 3783 (2a). Shank, P. & A. Molina R. 4610 (2b); 4696 (2b); 4719 (2a); 4766 (2b); 4783 (2a); 4850

(2b); 4966 (2b). Shattuck, O. 611 (3a); 1098 (3a). Skutch, A. 2660 (2a); 2890 (2a); 4371 (2a); 5201

(2c). Smith, D. et al. 13188 (1). Solis R., E 416 (2b); 495 (2a). Solomon, J. 5328 (2b); 13940 (1);

14104 (1). Soza, D. et al. 378 (2b). Standley, P. 41029 (3a). Starry, D. 28 (3a); 178 (3a). Stevens, W.

D,. 12153 (2b); 12493 (2b); 12836 (2b); 13083 (2b). Stevens, W. D. et al. 23470 (2a). Stier, F. 193

ae Sullivan, G. 58 (2a). Sytsma, K. 1523 (3a); 1975 (3a); 2650 (3a). Sytsma, K. & L. Andersson

1 (3b). Sytsma, K. & T. Antonio 2997 (2a). Sytsma, K. & W, D’Arcy 3610 (2a); 3690 (3a). Sytsma,

.et al. 2900 (3a); 4192 (3a).

abs, J. 18094 (2b).

Ule, E. 9641 (1). Utley, J. & K. Utley 1194 (2c); 3273 (2b); 3305 (2b).

Valerio, M., 651 (2c); 1268 (2a). Vasquez, R. et al. 5924 (1). Vineelli P, 252 (2b).

Wagner, M. 246 (2b), 623 (2b). Wallnéfer, B. 16-201287 (1); 18-8488 (1). Weberbauer, A. 1809

7 (2a); 21876 (2a); 21937 (2a). Webster, G. & G. Breckon 16516 (2a). Webster,
& R. Dressler 16745 (3a). Wedel, H. von 661 (2b); 932 (2b); 1109 (2b); 1536 (2b); 1769 (2b).

Wilbur, R. et al. 22713 (2a). Williams, L. et al. 29120 (2b). Woodson, R. & R. Schery, 906 (2b).

Woodson, R. et al. 1239 (2a); 1569 (3a); 1746 (2a).

Zuniga, R. et al. 392 (2a).

—

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UNA NUEVA ESPECIE DE POLIANTHES
(AGAVACEAE) DEL ESTADO DE
OAXACA, MEXICO

ELOY SOLANO CAMACHO

Herbario FEZA, Carrera de Biologia,
Facultad de Estudios Superiores Zaragoza, U.N.A.M.
A.P 9-020, Delegacién lztapalapa
09230 México, D.E, MEXICO

ABISAI GARCIA-MENDOZA

Jardin es Instituto de Biologia, U.N.A.M.
P 70-614, Delegacién ee
Ae DF, MEXICO

RESUMEN
Se describe e ilustra Polianthes bicolor, una nueva especie del estado de Oaxaca, Méxic
Esta especie muestra similitudes con Polianthes geminiflora (La Llave et Lex.) Rose var. pclae
Polianthes geminiflora (La Llave et Lex.) Rose var. clivicola McVaugh y Polianthes graminifolia
Rose.

ABSTRACT
peas bicolor from the state of Oaxaca, México, is described as new and illustrated.
his species is similar to Polianthes geminiflora (La Llave et Lex.) Rose var. ifl.
ee geminiflora (La Llave et Lex.) Rose var. clivicola McVaugh and Polianthes Ee oes
ose.

El género Polianthes es endémico de México y las especies que lo conforman
han sido poco estudiadas desde el punto de vista taxonémico. La ultima
revision fue hecha en 1903 por Rose, quien reconocié 12 especies. Durante
mucho tiempo, la delimitacidn de sus especies y de los géneros cercanamente
relacionados ha sido confusa. Actualmente se estd realizando la revisin sistematica
del género y las observaciones generadas durante el estudio, incluidas las
investigaciones anatomicas y la exploracién botdnica en toda su drea de
distribucién geografica, revelaron la existencia de una nueva especie que se
describe a continuacién.

pone bicolor Solano & Garcia-Mend., sp. nov. (Fig. 1). Tipo: MEXICO.
ah de Coixtlahuaca, en la base del Cerro Verde, 4 km al N de Marcos
eS y 8 km al S de Coixtlahuaca, pastizal, 2500 m, 7 Jul 1986, A. Garcta-Mendoza

et al. 2403 (HOLOTIPO: MEXU; ISOTIPOS: BRIT, FEZA, MEXU).

SIDA 18(2): 473-477. 1998

474 Sipa 18(2)

Herba perennis; lamina semisucculenta, plerumque undulata; bracteolae florum basalium
saepe in pedicello insertae; tubus floralis supra basim abrupte dilatatus; flores aurantiaco-
virides, lobis viridibus; filamenta in perianthii tubo 1.1—1.8 cm supra apicem ovarii inserta.

Hierba perenne con un bulbo de (2—)3—4(—5) cm de largo, (I—-)1.5-2.5
cm de didmetro; (3—)4—G6(—12) hojas basales de (5—)8—15 cm de largo, 0.6—
1(—1.4) cm de ancho, lanceoladas, lamina semi-suculenta, generalmente
ondulada; margen usualmente ondulado, papiloso, hialino; superficie lisa
a papilosa. Bracteas de la base de la goa ates 3-5(—7.5) cm de largo y
().2—0.5 cm de ancho, lanceoladas; infl de 24-40(-54) cm de longitud,
con 3—5(—9) nudos fértiles; flores geminadas, las inferiores a veces fusionadas.
Bracteas y bractéolas lanceoladas que disminuyen en tamano desde la base
de la inflorescencia hacia el 4pice, bractéolas de las flores basales frecuentemente
insertas sobre el pedicelo; pedicelos de 0.6—1.3 cm de largo, rojizos. Flores
de (2—)2.3—2.9 cm de largo, (1.6—)2.4—3(—5.5) mm de ancho en la base de
los I6bulos de la corola, tubo del perianto vere ensanchado por
arriba de la base; I6bulos externos de 2—3(—4) mm de largo, (1.7—)2—3(-4)
mm y en ocasiones de hasta 5 mm de ancho, de orbiculares a transversalmente
elipticos, apiculados, los internos con los apices redondeados y pilosos; color
de las flores anaranjado-verdoso, con los Iébulos verdes; estambres inclusos,
filamentos de 1.8—2.5(—2.8) cm de largo, insertos en el tubo del perianto,
1.1-1.8(—1.9) cm por arriba del dpice del ovario, filiformes, de color verde;
anteras de 4—5(~9) mm de largo, 1-1.5(-2.5) mm de ancho, oblongas, amarillo-
verdosas; estilo de (1—)1.9—2.6(—3.2) cm de largo; estigma trilobado. Fruto
semiesférico de 1.1 cm de largo por 1.1 cm de didmetro, Semillas de 4.5
mm de largo por 2.4 mm de ancho, de color negro, semicirculares vistas en
perfil, con uno de los lados recto.

Paratipos: MEXICO. Oaxaca: Municipio Teposcolula, 3 km al N de Yucunama, 24 Jul
1982, A. Garcta- bee 1052 (ENCB); Municipio Coixtlahuaca, Cerro Verde, S$ of
Coixtlahuaca, 7 Jul | , D. Frame et al. 317 (NY); Municipio de San Juan Mixtepec,
Cafiada Cerro . rea 20 km al NW de San Juan Mixtepec, 8 Jul 1988, /. Reyes 300
(MEXU); Municipio de San Juan Mixtepec, Yucu Shuun (Monte del Tesoro), 16 km al S
de San Juan Mixtepec, 6 Oct 1988, /. Reyes 881 (MEXU); Municipio de Teposcolula,
aproximadamente 5 km al NE de Marcos Pérez, 5 km al NE de Tierra Blanca, 3 km al NE
de Yodobada, a un costado del camino de terracerfa que va de lierra Blanca a Marcos Pérez,
26 Jul 1996, FE. Solano & Ma. del C. Lépez 857 (FEZA, MEXU); Municipio de Coixtlahuaca,
Cerro Verde al NE de Marcos Pérez, 7 Jul 1986, P Tenorio et al. 11656 (MEXU).

—

Polianthes bicolor se caracteriza por sus hojas semisuculentas, generalmente
con la l4mina y el margen ondulados, flores con el tubo del perianto abruptamente
ensanchado por arriba de la base, con Idbulos verdes y estambres insertos
sobre el tubo del perianto. Ademas, los estudios anatémicos, muestran la
ausencia de taninos en las células del mesofilo y de fibras en la vaina de los
haces vasculares de las hojas. Florece en los meses de julio a agosto; sus flores
no son fragantes, igual que en los otros taxa con los que aqui se compara.

SOLANO Y GaRcIA-MENDOZA, Una nueva especie de Polianthes 475

>

- os a
: He SPar2z2a—

Fic. 1. Polianthes bicolor. a) Planta con inflorescencia, b) Bulbo y cormo con raices contractiles,
c) Bractéolas insertas en los pedicelos, d) Flor completa, e) Flor disecada, f) Fruto, g) Semilla.

Ilustracién: a, c, d, e, de A. Garcia-M. 2403; b, f, g, de E. Solano C. & Ma. del C. Lépez B.
857.

476 Stipa 18(2)

Morfoldgicamente se relaciona con /? geminiflora (La Llave et Lex.) Rose
var. geminiflora, P geminiflora (La Llave et Lex.) Rose var. clivicola McVaugh
y P graminifolia Rose, de las que se distingue claramente por las caracteristicas
mostradas en la tabla 1.

Polianthes bicolor se desarrolla entre los 2300—2500 m, en pastizales, bosques
de Quercus y Pinus-Quercus con Agave, Anthericum, Opuntia, Salvia y Se-
dum, en suelos erosionados de color pardo a negro con textura arenosa y
material parental calizo.

La especie aqui descrita se localiza en la Mixteca Alta del estado de Oaxaca,
en la provincia fisiografica conocida como sistema montanoso del norte de
Oaxaca, en los distritos de Coixtlahuaca, Teposcolula y Juxtlahuaca.
Probablemente las plantas sefaladas por Howard (1986) para Huajuapan
de Leén, correspondan a esta nueva especie; el mismo autor se refiere a las
poblaciones en cuestién como raras y poco similares a Polianthes geminiflora,
ya que tienen flores de color escarlata, con Idbulos de la corola e interior
del tubo verdes.

Vasia 1. C isti folégi Polianthes bicolor, P geminiflora var. geminiflora,

ABLA L, g f
P geminiflora vax. clivicola y P graminifolia.

Cardacter Polianthes bicolor P. geminiflora P. geminiflora P. graminifolia
var. geminiflora var. clivicola
Lamina foliar Ondulada, rara vez Plana, rara vez Plana, Involuta, no
plana, cae a ada, no suculenta suculenta
culenta
‘Tubo del perianto Abruptamente ‘Tubular ‘Tubular Tubular

ensanchado por
arriba de la base

Color del tubo Anaranjado-verdoso — Anaranjado, Anaranjado, — Coral,
coral 0 rojol rojo 0 cora casi rosa 0 rojo
Color de los Iébulos Verde Anaranjado, Anaranjado Casi rosa

a veces verde

Insercién de 11-18 mm por 1.7—5 mm por 2—3mm por = 3.5—8 mm por
los filamentos arriba del ovario arriba del ovario — arriba del ovario arriba deli ovario
‘Taninos en las Ausentes Presentes Presentes Presentes

células del mesotilo

Fibras en la vaina Ausentes Presentes Presentes Presentes
del haz vascular

SOLANO Y GARCIA-MENDOZA, Una nueva especie de Polianthes 477

Polianthes bicolor es la especie con distribucién mas surefia del género,
sdlo conocida del estado de Oaxaca. En cuanto a los otros taxa, P geminiflora
(La Llave et Lex.) Rose var. geminiflora se distribuye del estado de Nayarit
a Tlaxcala y probablemente hasta el estado de Puebla, Polianthes geminiflora
(La Llave et Lex.) Rose var. clivicola McVaugh es endémica de Jalisco y Michoacan,
mientras que Polianthes graminifolia Rose solamente se ha registrado de
Aguascalientes y Jalisco.

El epiteto especifico hace referencia a la coloracién que presentan las flores.

AGRADECIMIENTOS

Este trabajo fue realizado con apoyo financiero de la CONABIO, a través
del proyecto FB291/H230/96. Asimismo, E. Solano agradece al CONACYT
la beca otorgada. Por otro lado, hacemos patente nuestro agradecimiento a
J. Rzedowski, T. Wendt, P. Davila y E Chiang, por la revisié6n cuidadosa
del manuscrito. F. Chiang preparé la traduccién al latin de la diagnosis y T.
Terrazas, asesor6 los estudios anatémicos. Nuestro reconocimiento a C. Correa
por su invaluable ayuda y compafia en la recolecta de especfmenes por toda
el drea de distribucién del género. A Elvia Esparza por la ilustracion realizada
y a los siguientes herbarios por el prest4mo de ejemplares: ENCB, FEZA,
MEXU y NY.

REFERENCIAS
Howarb, T.M. 1985. Stalking the Polianthes of Mexico-Part two, 1985. Herbertia 42:84—

87.
Rose, J.N. 1903. Studies of Mexican and Central American plants. Contr. U.S. Natl. Herb.

wa

Sipa 18(2

X
™
oO

BOOK REVIEW

Epwarb G. Voss. Michigan Flora. Part II]. Dicots (Pyrolaceae-Compositae).
1996. (ISBN 87737-040-0, hbk). Cranbrook Institute of Science Bulletin
61 and University of Michigan Herbarium, Ann Arbor, Michigan. Orders:
Cranbrook Institute of Science, 1221 N Woodward Ave., P.O. Box
MI 48303-0801. 810-645-3239 or 810-645-

illus-

801, Bloomfield Hills,
3203. $15.00 + shipping and handling. 622 pp, 771 maps, 267

peal

trations.

This ume is eee: alg and last of a series of volumes providing practical, user-friendly

keys, diagnc wracters, literature references and detailed ecological notes, supplemented

by 771 icituibution maps, 267 line drawings, 53 color photographs and keys to all the

vascular plant families occurring within the 86 counties comprising Michi per-

sonal experience with Part 1. Gymnosperms and Monocots, as an undergraduate student

at Michigan State, combined with my knowledge of Ed’s passion for detail and accuracy,
the previous two. The only item

tell me that this volume is an excelent reference as were
| consider missing from this fine book is that of a summary table, where total number of
habitats, number of families, genera, species, and perhaps endangered, threatened or rare
species, was included. While one can caution against such lists that often become restric-
tive and authoritarian, they are nonetheless, extremely useful to cc , land use
planners and resource managers, particularly those operating at the Gieex [ ales Regional
level. However, that is an insigifican blemish on an otherwise perfect publication. At
the incredibly low price of $15 this hard bound book on high quality paper ts a must

for every botanist’s bookshelf. on /. Pipoly Ill

Ilo All

Stipa 18(2): 478. 1998

POLLEN MORPHOLOGY OF THE GENUS
ECHINOPEPON (CUCURBITACEAE)

CONCEPCION RODRIGUEZ J. AND
RODOLFO PALACIOS-CHAVEZ

Departamento de Botdnica
Escuela Nacional de Ciencias Bioldgicas, I. PN.
Apdo. Postal 17-564
MEXICO, D.E CP.11340

ABSTRACT

Pollen descriptions of Echinopepon species and a palynological key are presented. Data
on apertures, exine, polarity, shape, and size of pollen grains are included. Pollen mor-
phology is Hees for ne species of Echinopepon re Echinocystis and in some Marah
species. The main features of each taxon are illustrated by photographs.

RESUMEN

Se presenta la descripcién del polen de las especies de Echinopepon. Se incluyen datos de
la apertura, exina, polarid: ad, forma y tamafi granos, asi como una clave palinolégica
para su identificacién. La mor fologia polinica es Cieeercicien apres y distintiva
de Echinocystis y de algunas especies de Marah. Se ilustra con fotografias las caracteristicas

principales de cada taxon

Echinopepon is an American genus, whose distribution extends from
southeastern United States to Argentina. It has seven species (Rodriguez,
in press): E. cirrhopedunculatus Rose, E. coulteri (A. Gray) Rose, E. gemellus
(Ser.) Rodriguez, E. minimus (Kellogg) S. Watson, E. pee (Benth.)
Rose, £. racemosus (Steud.) C. Jeffrey, and EF. wrightii (A. Gray) S. Watson
(Table 1); the first four are endemic to Mexico. All species are herbaceous,
generally with an annual life cycle; have characteristic capsular, echinate,
rostrate, and operculate fruit; and white or white-green pentamerous flow-
ers, five stamens with filaments joined and anthers forming heads.

Jeffrey (1978) placed Echinopepon in the tribe Cyclanthereae. He later
(Jeffrey 1990) placed it in tribe Sicyeae subtribe Cyclantherinae, showing
its close relationship to the genera Marah and Echinocystis as demonstrated
by the large number of morphologic characteristics which it shares with
these genera. They are distinguished from Echinopepon by two or three sta-
mens and globose fruits with apical and irregular dehiscence. Marah is dis-
tinguished also by its perennial habit and swollen seeds.

Palynologically the tribe Cyclanthereae (Jeffrey 1964) is characterized

Stipa 18(2): 479-491, 1998

E. minimus (Kellogg) S. Watson

E. pubescens (Benth.) Rose

E:

E.

E. gemellus (Ser.) ¢

480

was described as having multizonocolporate pollen grains.

The pollen morphology of Echinopepon species has been little studied.
found similarities between Apatzingania arachoidea Di-
eter. pollen grains (asymetrical polycolpate, spinulose and exquisitely fo-

Dieterele (1974) f

SipA 18(2)
by 4-8-colporate and punctitegillate pollen grains (Jeffrey 1964), the same
as the subtribe Cyclantherinae (Jeffrey 1990). This tribe was also described
by Marticorena (1963) as having 4 to polycolporate pollen grains except,
Echinopepon wrightii (A. Gray) S. Watson (as Echinocystis wrightii A. Gray)
which has only polycolporate pollen grains. In a palynologicalal classifica-
tion of Cucurbitaceae from India (Shridar & Sing 1990), subtribe Cyclantherinae

veolate-reticulate) and those of Echinopepon cirrhopedunculatus and E. ge

TABLE 1. [he species of Echinopepon.

Accepted name

E. cirrhopedunculatus Rose
E. coulteri (A. Gray) Rose

i
>, Rodriguez :

Echinoc ystts ai . ay) Cogn.,
1897;

tA

PE PSS ey

Synonyms

1877;
Echinopepon confusu

us
‘chinopepon a hse Roce 1897,

fe rinopepon milleflorus Naudin, 1866;

; ad on Bis eae omer 1866;
s gemella

WNOCYSE (Ser.) € , 1877;
‘hinocystis a ua (Nandi) c ogn., 1877;
ch TNO oe sts lea er.) C 1877
NOC) Or qué

SS

oe brevspina Cogs 1881;

Ee ra ei ae (Ser.) Rose, 1897.

fx

7 ‘
C. Jeffrey -

racemosus (Steud.)

a Nig minima (S. Watson) on

Ec. hinopepot

ee Bele oo minimt as OK Kell.) S. Watson var, ror iedlnnle S. Gentry
955.

pine “ystis Us patie Cogn.,

ing
peninsular H.S. Ge

tocking, 1

1877;
hinocystis pubescens (Benth.) Cogn., 1877;

ee Aasbiandis (Cogn, ) Rose, 1897.

inopepon ee Dee 1866;

inocystis lanata

- innosth pal rpa e. ogn., oe
Echinocystis murtcata C ae 1878;
Echinocystis araneosa Griseb., 18

ieshe

Ec
a

wrightit (A. Gray) S. Watson

79;
schinocystis macroc os aot 890;

inopepon ja iscanus Rose, 1897
inopepon lanatus (Cogn.) Rose, 1897;
ypepon p / Rose, 1897

moc ystts glutinosa rae
inocystts lon gispina Co

hinocyst is panteulata C og 18
chinocystis wrightii (A
; hid

ae
Cray) Cogn., 1877;
iain ‘ogn.) Rose, 1897;
inopepon nelsoni I , 186
inopepon laeoe (Cc ogn. ) J.V. Dieterle, 1976.

hinopepon racemosus (Steud. ) Mista Crov., 1955.

)

Ropricurz AND PaLacios-CHAvez, Pollen morphology of Echinopepon 48]

mellus (as Echinocystis gemella). Dieterle (1976) also found that different
types of apertures in pollen grains seemed to be associated with thecae form,
replicate thecae occurring with non-colpate grains and U-shaped thecae occurring
with multicolpate grains. Stafford and Sutton (1992, 1994) indicated that
certain species of Echinopepon had pantozonocolporate pollen grains, cor-
roborating Dieterle’s idea about palynological similarities between the genera
Apatzingania and Echinopepon, however, there are enough macromorphological
differences to distinguish Apatzingania from Echinopepon. Palacios-Chavez
et al. (1995) described Echinopepon coulteri pollen the valley of Mexico as
7-colpate, perreticulate and suboblate grains and F£. gemellus pollen (as
Echinopepon milleflorus) as 14-colpate, punctitegillate and suboblate.

MATERIALS AND METHODS

Pollen samples of the seven taxa of Echinopepon and the relatives Echinocystis
and Marah were taken from herbarium specimens at: Escuela Nacional de
Ciencias Bioldgicas, IPN, México (ENCB); Herbario Nacional of México,
UNAM, México (MEXU); Instituto de Ecologia, Bajfo, México (IEB) and
Field Museum (F). The samples were prepared using the standard acetoly-
sis method of Erdtman (1943) and mounted in glycerine jelly. The samples
were examined by the light microscope. The slides are deposited in the pollen
collections of the Palynology Laboratory, Department of Botany, ENCB,
Mexico.

RESULTS

a

Palynological key to the species Echinopepon

1. Grains stephanocolpate, 7—16-colpate or 7—17-colporoidate, reticulate or

punctitegillate.
2. Grains 7-colpate or colporoidate.
3. Grains 7-colpate, reticulate E. coulteri (Figs. 6-11)
3. Grains 7-colporoidate, punctitegilate..4 3 ee cc.:. E. pubescens (Figs. 22-27)
2. Grains 14—16-colpate or 9-10-colporoidate.
4. Grains 9-10-colporoidate E. minimus (Figs. 16-21)
4. Grains 14—16-colpate.
5. Grains 14-colpate, punctitegillate E. gemellus (Figs 12-15)
5. Grains 16-colpate, reticulateseent.s2..:s..4.. E. cirrhopedunculatus (Figs. 1—5)

1. Grains 10—12-perirrugate or 7—8-pericolpate, punctitegillate-echinulate or
perreticulate.
6. Grains perirrugate, 10-12 rugas, echinulate E. racemosus (Figs. 28-34)
6. Grains pericolpate, perreticulate E. wrightii (Figs. 35-41)

Descriptions of pollen grains

Echinopep hoped latus Rose (Figs. 1-5). MEXICO. MORELOS: Zacatepec
C. Rodriguez et al. 1856 (ENCB).

482 Stipa 18(2)
Grains usually stephanocolpate, 16-colpate (sometimes with 15 or 17

colpi), semitectate, spheroidal, (88—)72—104 x 77—104(—82) um P/E 1.07.

Polar view circular, (103—)92—114 pm in diameter. Exine 5.6 pm, thick,

sexine 4 pm and nexine 1.6 pm, thick reticulate, with colpus thin; mem-

brane smooth.

Echinopepon coulteri (A. Gray) Rose Figs : a MEXICO. HIDALGO: Cerro
Gordo, 5 km W of Pachuca, /, Rzedowski 3352: iD),

Grains usually stephanocolpate, 7-colpate, ae 8-colpate, semitectate,
suboblate, (104—)83-117 x 87-142(—126) pm. P/E 0.82. Polar view cir-
cular (1 18—)77—154 pm in diameter. Exine 4 pm, thick sexine 3.2 pm and
nexine 0.8 pam, thick, reticulate superficially, colpus shallow, membrane smooth.
Echinopepon gemellus (Ser.) C. Rodriguez (Figs. 12-15). MEXICO. VERACRUZ:

Camino a Tatatila, Las Vigas, F Ventura 17716 (ENCB).

ani

Grains stephanocolpate, 14-colpate,tectate, suboblate, (76—)66—97 x 66—
106(—90) pm. P/E 0.84. Polar view circular (88—)70—112 tm in diameter.
Exine 4 pm, thick, sexine 3.2 um and nexine 0.8 pm, thick, punctitegillate
superficially, colpus thin; membrane smooth.

Echinopepon minimus (Kellogg) S. Watson (Figs. 16-21). MEXICO. Baja

CALIFORNIA: Sierra de Plateros, Mulege, D. Breedlove 60907 (ENCB).

Grains usually stephanocolporoidate, 10-colporoidate, sometimes 9-
colporoidate, semitectate, spheroidal, (7 1—)60—85.6 x 74—84(—80) pm. P/
E 0.95. Polar view circular, (85—)80—96 tim in diameter. Exine 5 um, thick,
sexine 3.2 um and nexine 1.8 pm, thick, reticulate superficially, colpus with
smooth membrane. Pores diffuse.

Echinopepon pubescens (Benth.) Rose (Figs. 22-27). MEXICO. Oaxaca:

Chilapa, Tamazulapa, /, Rzedowski 34498 (ENCB).

Grains stephanocolporoidate,7-colporoidate some times 6-colporoidate,
tectate, spheroidal, (94—)81—110 x 81—103(—94) um. P/E 1.0. Polar view
circular (102—)91—112 pm in diameter. Exine 4 ym, thick, sexine as thick
as sexine, punctitegillate superficially, thin colpus, smooth membrane, bordered
with margin 2—3 pm wide.

Echinopepon racemosus (Steud.) C. Jeffrey (Figs. 28-34). MEXICO. Oaxaca:

Ojitlin, Buena Vista, F Ventura 15539 (ENCB).

Grains perirrugate, tectate, apolar, spheroidal, (94—)81-110 x 91-112(-
102) pm in me Exine 5.6 im, thick, sexine as thick as nexine, echinulate
superficially. Rugas 10 or 12 of 16 pm long, situated superficially, with verrucate
membranes. Some grains have a triradiate colpus.

Echinopepon wrightii (A. Gray) S. Watson (Figs. 35-41). MEXICO, MICHOACAN:

3 km NW of Eréndira Cardcuaro, /. Reedowski 35510 (ENCB).

Grains stephanocolpate, usually 7-colpate,sometimes 8-colpate, semitectate,

RoprIiGuez AND Patacios-CHAVEZ, Pollen morphology of Echinopepon 483

Fics. 1-11 —Echinopepon i ee Big. 1 Equatorial view; Fig. 2. Polar view;
Fig. 3. Optical section exine (x 1000); Fig. 4. Equatorial view (x 800); Fig. 5. Polar view
(x 800). ae coulteri. Fic. 6. Tae section exine; Fig. 7. Polar view surface x
800); Fig. 8. Polar view optical section (x 800); Fig. 9. ee view surface (x 800);
Fig. 10. Reticulate ee Fig. 11. Optical section (x 1000

SipA 18(2)

Fics. 12-22. —Echinopepon gemellus. Fig. 12. Equatorial view (x 800); Fig. 13. Polar view
(x 800); Fig. 14. Polar view, oe ae Me 15. Optical section.(x 1000). Hchinopepon
minimus. Fig. 16. Equatorial view (x 800); Fi ig. 17. Equatorial view, reticulate seus Fig.
18. Optical section (x 1000); Fig. 19. oe view (x 800); Fig Ig. 20. E ‘ichinulate apocalpium;

Fig. 21. Optical section (x 1000). Fig. 22. £. pubescens. Equatorial view (x 800

RopriGuez AND Patacios-CHAvez, Pollen morphology of Echinopepon 485

Fics, 23-31. —Echinopepon pubescens. Fig. 23. Punctitegillate surface; Fig. 24. Polar view
(x 1000); Fig. 25. Polar view, optical section (x 800); Fig. 26. Colpi and punctitegillate
es Fig. 27. Exine thick. ate 28-31. Echinopepon racemosus. (x 800). Fig, 28. Perirrugate;

29. Ruga aequatorialis; Fig. 30.Ruga and echinulate surface; Fig. 31. Echinulate sur-

ae

486 SIDA 18(2)
spheroidal, (81—)71—90 x 86-94(—89) pm. P/E 0.94. Polar view circular,

(94-)80—-108 um in diameter. Exine 5 um, thick, sexine and nexine with
the same thickness, perreticulate superficially, 7 colpi in equatorial posi-
tion but 8-colpate where one colpus ts over a pole.
ee lobata (Michx. eee : A. Gray (Figs. 42-49). U.S.A. OHIO.
DARKE CO.: M.A. Vincent 1372 (MEX
Grains stephanocolporate, ay ee often 6-colporate, tectate,
suboblate, (G0—)53—68 x 64-77(—70) um. P/E 0.85. Polar view pentago-
nal or circular of (69-)64—72 um in diameter. Exine 4.8 um, thick, sexine
and nexine the same thickness, punctitegillate. Colpi with scabrate mem-
branes, margocolpate. Pores lalongate, (17—)12—20 pm x (7.6-)4—9 um,
with granulate opercula.

Marah gilensis (Greene) Greene (Figs. 50-53). U.S.A. ARIZONA: 1968, P Stewart
sm. (MEXU).

Grains stephanocolporate, usually 4-colporate, often 5-colporate, semitectate
spheroidal, (60—)56—-63 x 47-60(—56) pm. P/E 1.07. Polar view quadran-
gular, (56—)53-63 pm in diameter. Exine 4 pm, thick, sexine thicker than
nexine, reticulate (clava forming a reticulum). Colpi with smooth mem-
branes, acute ends, with broad margin 3 um wide. Transversal colpi (18-
)14—21 x 3-5(—4) pm, with scabrate opercula.

Marah macrocarpa (Greene) Greene (Figs. 54—57). MEXICO. BAJA CALIFORNIA:

R. Moran 29109 (ENCB).

Grains stephanocolporate, 4-colporate, semitectate, subprolate, (69-)65—
76 Xx 50-62(-57) um. P/E 1.2. Polar view circular (64—)50—-73 pm in di-
ameter. Exine 3 pm, thick, sexine and nexine the same thickness retipilate.
Colpi with acute ends, with broad margin 3 um wide, syncolpate at a pole,
membranes smooth. Circular pores (7—)5—10 jm in diameter.

DISCUSSION

Pollen morphology i is characteristic for Echinopepon and Echinocystis and
for some species of Marah.

Pollen from Echinopepon species was found to be variable between spe-
cies, especially in the number of openings and in their ornamentation. The
highest number of colpi was sixteen (Table 2) and contrasts with Jeffrey’s
data (4—8-colporate) for tribe Cyclanthereae (1964) and subtribe Cyclantherinae
(Jeffrey 1990).

The pollen ornamentation was found to be reticulate (R), echinate (E),
perreticulate (PR) and punctitegillate (PT) in different species (Table 2).
The punctitegillate type was reported by Jeffrey (1964,1990) and Stafford
and Sutton (1994) for Echinopepon coulteri, E. racemosus and Echinocystis
lobata.

RopriGuez AND Patacios-CHAvez, Pollen morphology of Echinopepon 487

rl

Fics. 32—41.—Echinopepon racemosus. Fig. 32. Echinulate surface; Figs. 33, 34. Triradi-
ate colpi (x 1000). Echinopepon wrightii. Fig. 35. Equatorial view (x 800); Fig. 36. Polar
view (x 800). 37. Perreticulate surface; Fig. 38. Equatorial view; Fig. 39. Polar view, op-
tical section (x 800); Fig. 40. Colpus over a pole; Fig. 41. Perreticulate surface (x 1000).

Sipa 18(2)

488

Fics. 42—49.—Echinocystis lobata. Fig. 42. Equatorial view, punctitegillate surface; Fig. 43.
Optical section; Fig. 44. Polar view (x 1000); Figs. 45-49. Optical section (x 800).

RopriGurz AND PaLacios-CHAveEz, Pollen morphology of Echinopepon 489

Fics. 50-57.—Marah gilensis. Fig. 50. Equatorial view; Fig. 51. Optical section; Fig. 52
Polar view surface; Fig. 53. Optical section (x 1000); M. macrocarpa. Fig. 54 Howton
view surface (x 1000); Fig. 55. Syncolpate grain; Fig. 56. Polar view syncolpate (x 800);
Fig. 57. Optical section (x 1000).

490 Sipa 18(2)

TABLE 2. Palynological characteristics Echinopepon species.

P (um) E (um) P/E Ornam. N° Colpi Exine Thecae
x xX ‘Type
E. cirrhopedunculatus 88 82 1.07 R 16 5.6 U
E. coultert 104 126 0.82 R 7 4.0 U
E. gemellus 76 90 0.84 PT 14 4.0 U
E. minimus 71 80 0.95 R 10 por, 5.0 U
E. pubescens 94 94 1.00 PT 7 por. 4.0 Replicate
E. racemosus 94 102 E 5.6 Replicate
E. wrightii 81 89 0.94 PR 7 5.0 Straight
or
Arched

P. Polar Axis E. Equatorial Axis P/E. Relation between polar axis and equatorial axis.

Echinopepon cirrhopedunculatus, E. coulteri, E. gemellus and E. minimus
have U-shaped thecae and colpate or colporoidate pollen grains, while F.
racemosus has replicate thecae and its pollen grains lack longitudinal colpi.
E. pubescens has replicate thecae and colporoidate pollen grains, while the
thecae of E. wrightii are straight or arched with a 7-8 colpate pollen. The
results presented in Table 2 agree with those reported by Dieterle (1976)
for E. gemellus (as E. torquatus), E. racemosus (as E. horridus) and E. wrightii
(as E.paniculatum).

The tectate pollen grains and the absence of longitudinal colpi observed
in E. racemosus may suggest that it is the most advanced species of the ge-
nus. [his taxon is quite variable regarding the shape of its leaves, number
of tendrils and branches, degree of pubescence and wide geographical dis-
tribution from Mexico to Argentina. On the other hand, F. cirrhopedunculatus,
E. gemellus and E. minimus may be considered less advanced because their
pollen grains are semitectate and 10-17 colpate-colporoidate; they are re-
ported as endemic to Mexico (Rodriguez 1995). Although E. pubescens is
also considered endemic to México, it has 6—7-colporoidate pollen grains.

In conclusion, the pollen of Echinopepon, Echinocystis and Marah is of-
ten distinctive, which in some cases helps in taxonomic determination.
Echinopepon pollen characters have diagnostic value in identification of the
seven accepted species.

REFERENCES
Dievrerce, J.V.A. 1974. A new geocarpic genus from Mexico: Apatzingania (Cucurbitaceae)
Brittonia 26:129-132.
1976. Flora of Guatemala. Cucurbitaceae. Fieldiana Bot. 24, XI(4):306-395.
ERDTMAN, G. 1943. An introduction to pollen analysis. The Ronald Press Co., New York.
Jerrrey, C. 1964. A note on pollen morphology in Cucurbitaceae. Kew Bull. 17:473-477.

RopriGuez AND PaLacios-CHAvez, Pollen morphology of Echinopepon 49]
. 1978. Further notes on Cucurbitaceae IV. Some new world taxa. Kew Bull. 33:347-
380

. 1990. Appendix: An outline classification of the Cucurbitaceae. In: D.M. Bates,
R.W. Robinson, and C. Jeffrey, eds. Biology and utilization of Cucurbitaceae. Cornell
Univ. Press. Ithaca, New York. Pp. 449-463

Marticorena, C. 1963. Material para una ee de la morfologfa de polen de
Cucurbitaceae. Grana ae 4:78-9

Patacios-Cu.R., Ma.L. ARREGUIN S., y D.L. ce G. 1995. Morfologia de los sen
de as de la familia ae del Valle de México. Anales Esc. Nac. Ci. Biol.
40:1

ee Ze < C. 1995. Distribucién aes del género papa (Cucurbitaceae).
Anales Inst. Biol. Univ. Nac. México. Ser. Bot. Z 1-18

. Taxonomia del género einen (In pres

SHRIDHAR and D. SING. 1990. Palynology of the Tadian Cucurbitaceae. In: D.M. Bates,
R.W. Robinson and C. Jeffrey, ia ee and utilization of Cucurbitaceae. Cornell
Univ.Press. Ithaca, New York. Pp. 208.

STAFFORD, PJ. and D.A. SUTTON. oF ee morphology of the Cyclantherinae. C. Jeffr.
(Tribe Syceae Schrad. Cucurbitaceae) and its taxonomic significance. 8" Internat. Palynol.
Congress, Aix en Provence. Abstracts. P. 140.

. 1994, Pollen morphology of the Cyclantherinae C. Jeffrey (Tribe ae ae

(Gaines and its taxonomic significance. Acta Bot. Gallica 141:171-

S

Sipa 18(2)
BOOK REVIEW

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This book contains practical keys, shortcut identification tables, and descriptions for 299

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with salient features of closely related or similar-looking species, and a fold. -out grass identification
e. A vegetative key runs from page 9 to page 17, and while not dichotomous, seems

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Pipoly II

IDA 18(2): 492. 1998

VARIATION IN THE BERLANDIERA PUMILA
(ASTERACEAE) COMPLEX

G.L. NESOM
BONAP-NCBG
Coker Hall CB 3280
University of North Carolina
Chapel Hill, NC 27599-3280, U.S.A

B.L. TURNER
Department of Botany
University of Texas
Austin, TX 78713, U.S.A.
ABSTRACT
Plants from southeast Ti dadj I iously identified in part as Berlandiera

xbetonicifolia are regarded here as seogtaphic. digaaeeee pumila and are treated as B.
pumila var. dele Nesom & Turner, var. nov. Variety pumila and var. <—o differ in
features of cauline and foliar vestiture, that of one pee i: af surfaces allow
consistent separation. [he type collection of B. ¢ ia (Hook.) Small (Silphinm . Dinh lien
Hook., 1835) represents a species distinct Fane B ‘i ila sa comprises plants aes
identified by the name B. texana DC. (1836), which is displaced by the former. While
morphology of B. pumila var. scabrella has suggested that it originated as a hybrid ean
typical B. pumila and B. betonicifolia, there is no evidence of current hybridization be-
tween these two species.

Key worbs: Berlandiera, B. pumila, B. betonicifolia, Asteraceae

RESUMEN

Plantas del sudeste de Texas y la parte adyacente de Louisiana identificadas a
en parte como Berlandiera xbetonicifolia son consideradas aqui como variantes geogra
de B. pumila y son tratadas como B. pumila var. scabrella Nesom & Turner, var. nov. ae
variedad pumila y var. scabrella difieren en caracteristicas del indumento caulinar y flee
siendo la del haz la que permite una mejor separacidn. El tipo de B. betonicifolia (Hook.)
Small (Siphinm betonicifolium Hook., (1835) ese una especie distinta de B. pumila
y comprende Pp B. texana DC. ( 836), que es desplazada
por la anterior. Mientras que la mor an de B. pumila var. scabrella ha sugerido que se
formé como un hibrido B. pumila tipica y B. betonicifolia, no hay evidencia de hibridacién
actual entre estas dos especies.

Shinners (1951) followed Small (1903, 1933) in treating Berlandiera pumila

(Michx.) Nutt. as a species restricted to the southeastern United States east

SpA 18(2): 493-502. 1998

494 Sipa 18(2)

of the state of Mississippi; its vicariad west of the Mississippi River was identified
as B. dealbata (Torr. & A. Gray) Small. Turner and Johnston (1956) re-
turned to the concept of Torrey and Gray by including the western plants
within a more broadly defined B. pumila, acknowledging the existence of a
distributional hiatus between the two geographic segments of the species.
The western segment of B. pumila occurs primarily in east Texas and adja-
cent Louisiana (Fig. 1), barely reaching into the adjacent corners of south-
eastern Oklahoma and southwestern Arkansas.

Shinners (1951) accepted the existence of Berlandiera betonicifolia (Hook.)
Small as a distinct species similar to western B. pumila (B. dealbata in his
sense) in morphology and geography. Turner and Johnston (1956) viewed
B. betonicifolia as intergrading with both B. pumila and B. texana DC. and
adopted the earlier combination B. texana var. betonicifolia (Hook.) Torr.
& A. Gray. This group of entities/taxa is referred to in the present study as

seeteeeecu Man
yA on a I
RO a ee
ea
WIL OE ai NN
Raa wat |

— 2Os Berlandiera pumila

Fi ns var. pumila (white form) O
HY

var. pumila (green form) @

var. scabrella ®

Fic. 1, Distribution by county of Berlandiera pumila in Texas, Louisiana, Arkansas, and

Oklahoma.

NESOM AND TurNER, The Berlandiera pumila complex 495

the “B. pumila complex.”

In a biosystematic study of the whole genus, Pinkava (1967) found that
crosses between all combinations of species of Berlandiera were easily made
under greenhouse conditions and he provided evidence of natural hybrid-
ization between many of the taxa. All entities of the genus are reported to
have an apparently invariant chromosome number of =15 (see Pinkava
1970 for original counts and summary of previous literature). Based on
the morphology of artificially constructed hybrids, Pinkava observed that
“although the [GH] type of B. betonicifolia lies between the F; [of B. pumila
and B. texana] and B. texana, it is definitely an intermediate of the two
species and its name has been retained for the hybrid group” (p. 292). Pinkava
used the designation “B. xéetonicifolia” and has been followed in this by
recent treatments that have adopted his view.

Pinkava interpreted the Berlandiera pumila complex in east Texas (his
Fig. 4, p. 294) essentially as an intergrading nexus between B. pumila, B
betonicifolia, and B. texana. In B. betonicifolia, “most characters are inter-
mediate to those of the putative parents in varying degrees and combina-
tions over its range of east-central Texas and western Louisiana” (Pinkava
1970, p. 1625). He found apparent intermediates between B. texana and
B. betonicifolia, however, to be rare. Although his histogram of hybrid in-
dex values in the “Western B. pumila/texana complex” (Fig. 2b, p. 290)
indicates that he observed several populations of such intermediates, only
one such collection is mapped on his Fig. 4 (1967); it is in central Louisi-
ana (Rapides Parish), where B. pumila apparently is absent but where B.
texana (sensu Pinkava) is known by a number of collections (Fig. 2).

In the Louisiana flora, Gandhi and Thomas (1989) and MacRoberts (1989)
recognized both Berlandiera pumila and B. xbetonicifolia, showing these
two taxa with nearly identical geographic ranges. MacRoberts (1989, p
313) noted that the latter “seems to be inordinately common compared
with its putative parents.” Berlandiera pumila and B. xbetonicifolia both
have been recognized from the southeastern corner of Oklahoma (Taylor
and Taylor 1994).

Berlandiera xbetonicifolia (sensu Pinkava) was reported from the south-
western corner of Arkansas (Miller Co.) by Orzell and Bridges (1987, p.
88), who adopted Pinkava’s 1 interpretation of variability among the plants
they observed. They noted that “Our specimens mostly fit the put ackcross
hybrid of Berlandiera xbetonicifolia with B. pumila, . . although some of
our specimens have dense, matted white-tomentose pubescence, and could
be referred to B. pumila.” Smith (1994) identified these Arkansas plants
simply as B. pumila.

Cronquist (1980) did not include Berlandiera betonicifolia among the
three species formally recognized in his treatment of the genus for the southeastern

496 Sipa 18(2)

Fic. 2. Distribution by county of Berlandiera betonicifolia (sce text for definition) in Texas
and Louisiana. The species also occurs northward through Oklahoma into northern Ar-
kansas, Missouri, and southern Kansas.

United States. He referred to it in introductory comments simply as the
name covering hybrids between B. pumila and B. texana.

Despite the considerable previous attention given to patterns of varia-
tion and accompanying taxonomy of plants of the Berlandiera pumila complex
in Texas and Louisiana, field experience in east Texas and adjacent Louisi-
ana suggested that the pattern of variation in this group a be viewed
from another perspective. [his problem was approached in more detail eiees
study of numerous collections available in regional herbaria and field 0
servations in 1996 and 1997 covering Anderson, Angelina, Cherokee, Har a
Houston, Jasper, Montgomery, Nacogdoches, Newton, Polk, Rusk, Sabine,
San Augustine, San Jacinto, Shelby, Trinity, Tyler, Upshur, Van Zandt, Walker,

and Wood counties, Texas, and Vernon Parish, Louisiana.

THE IDENTITY OF BERLANDIERA BETONICIFOLIA
Pinkava did not examine the original type material of Si/phium betonicifolium,
but based on a photo, he noted (1967, p. 297) that “Drummond’s holo-
type [of S. betonicifolium] more closely resembles B. texana {than B. pumila]
and was assigned to it [B. texana] as a variety by Torrey & Gray (1842),

Nesom AND Turner, The Berlandiera pumila complex 497

who commented that other specimens fitted neither varietal description
completely.” Gray annotated at least two of the Drummond sheets now at
K (see below) as “Berlandiera texana DC. 8 betonicifolia Torr. & Gray.” The
present examination of the original material at K shows it to be the species
that has long been identified as Berlandiera texana DC., which de Candolle
named a year later than Hooker's S. betonicifolium.

The material at K consists of four sheets, which are similar among themselves
in habit and other morphological features, and it seems likely that Drummond
collected all of this material at the same site. The lectotype (designated below)
bears detailed sketches of the ray and disc achenes, which were probably
used by Bentham in his preparation of the Compositae treatment for Gen-
era Plantarum.

The correspondingly modified nomenclature for this species is summa-
rized below.

Berlandiera betonicifolia (Hook.) Small, Fl. SE U.S. 1246, 1340. 1903.
Silphium betonicifolium Hook., Comp. Bot. Mag. 1:99. 1835. Berlandiera texana
var. betonicifolia (Hook.) Torr. & A. Gray, Fl. N. Amer. 2:281. 1842. Berlandiera
xbetonicifolia ( (Hook.) Small sensu Pinkava, Brittonia 19: 297. 1967. Type: (as cited
in the protologue and on the label, see comments in text): [United States, Louisi-
ana], New Orleans, 1833, 7) Drummond s.n. (LECTOTYPE, designated here: K};
ISOLECTOTYPES: K-3 sheets!; probable ISOLECTOTYPE: “CHI. Geer photos of two of
the K isolectotypes are on file at TE
Berlandiera texana DC., Prodr. 5:317. 1836. Tyre: TEXAS: between the Trinity River
and Bejar {San Antonio], Jun 1828, /.L. Berlandier 2044 (HOLOTYPE: G fiche!; IsoTYPES:
P)

On the lectotype of Silphium betonicifolium (and one of the isolectotypes),
presumably written by Hooker himself, the collection locality and date are
given as N. Orl. 1833.” The possible GH type material of S. betonicifolium

was annotated by B.L. Turner and M.C. Johnston in 1955: “Possible isotype,
as questioned by Gray (Fl. N. Amer. 2:281. First edition). Drummond’s
plant possibly came from Texas—‘New Orleans’ being a citation error.” Gray
himself (1842) cited the locality of the type collection as “New Orleans?,”
apparently questioning its provenance. There are no other historical or re-
cent collections of any species of Berlandiera from southeastern Louisiana
and none from Mississippi, and it seems unlikely on this basis that Drummond
made the “betonicifolia” type collection in the area of New Orleans.

One of the isolectotypes bears a glued annotation label (pasted over the
lower stem), presumably in Asa Gray’s script: Berlandiera texana DC. Torr.
Gr. Fl. N. Am. 2.286.” At the bottom of the label, in print, there is a line
“Jacksonville (Amer. Bor.) Drummond 1833.” Gray (presumably) crossed
out the word Jacksonville and wrote above it “Louisiana.” An early Texas
site for “Jacksonville” is not known to the present authors.

498 Sipa 18(2)

Drummond sailed from New Orleans to reach Texas (Brazoria County)
in March 1833, and during his 21-month stay in Texas (Geiser 1948), he
stayed or traveled in areas where Berlandiera pumila var. pumila occurs (e.g.,
Fayette, Austin, Lee, Colorado, Fort Bend counties), but he did not travel
in the area where plants previously identified as B. xbetonicifolia (see be-
low) occur (Fig. 1). Geiser’s account shows that Drummond passed through
at least one county (Gonzales Co.) where B. betonicifolia (of the present
interpretation) is known to occur (Fig. 2). From among the 100 or more
collections of the Berlandiera pumila complex at LL, VEX, the plant most
similar to the Kew types is a collection (Gentry & Smith 2024, LL) from
Victoria Co., which is only a short distance from Gonzales County.

Considering the information presented in the present study, Drummond
probably collected the plants of the type material in south-central Texas at
some locality near the Guadalupe River, which traverses both Gonzales and
Victoria counties. Drummond collected along this river (fide Gaiser) be-
tween 24 April and 26 September, 1834, at which time he was assembling
his “set II.” One of the isolectotypes bears the handwritten notation “Texas
II, no No. [s.n.], Drummond.”

With his last set of collections, Drummond sailed from New Orleans in
December 1834 on his return to Europe. He disappeared enroute (appar-
ently in Cuba) but his specimens arrived safely.

THE NATURE OF BERLANDIERA BETONICIFOLIA

Berlandiera betonicifolia (B. texana of previous interpretations) occurs from
panhandle and south-central Texas and adjacent Louisiana (Fig. 2) north-
ward through Oklahoma into northern Arkansas, Missouri, and southern
Kansas (Pinkava 1967; Turner and Johnston 1956; Great Plains Flora Assoc.
1986). This species can be separated from the B. pumila complex without
intergrades by making comparisons in the key below. Collections of both
B. betonicifolia and B. pumila have been made in the same counties in cen-
tral and east Texas (Victoria, Bastrop, Burleson, Gonzales, Travis, Fayette,
and Shelby) and parishes of adjacent Louisiana (Caddo and DeSoto) (Figs.
1 and 2), but in this region of geographic overlap, there are no plants sug-
gestive of hybridization and/or introgression between them. For the most
part, plants that previously have been interpreted as such hybrids (B. pumila
var. scabrella, as identified here) occur outside of the present geographic
range of B. betonicifolia.

As noted by Pinkava (1967), the lower stems of plants of Berlandiera
betonicifolia from parts of Texas tend to be silky-pilose with soft, white hairs
while those from further north have shorter, thicker, and slightly vitreous
hairs. Infraspecific variation in leaf shape and margin also exists within the
species, but this variability apparently does not bear on its relationship to
B. pumila.

Nesom AND TurNeR, The Berlandiera pumila complex 499

Based primarily on perceived similarities in stem leafiness and leaf inser-
tion, Turner and Johnston (1956) regarded Berlandiera betonicifolia (as B.
texana) and B. pumila var. scabrella (as B. betonicifolia) as conspecific and
intergrading. [he current study, in contrast, concludes that B. betonicifolia
is distinct from other closely related entities.

VARIATION WITHIN BERLANDIERA PUMILA

Plants from east Texas and western Louisiana previously identified as
Berlandiera pumila and B. xbetonicifolia are best represented as two enti-
ties within a single species (B. pumila), differing in features of vestiture.
The nomenclatural summary for typical B. pumila and the newly recog-
nized variant are given here.

Berlandiera pumila (Michx.) Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:342.
1841. Silphium pumilum Michx., Fl. Bor.-Amer. 2:146. 1803. Type: UNITED STATES.
FLoripa: [no date], A. Michaux 10 (HOLOTYPE: P fiche!).

Berlandiera tomentosa var. dealbata Torr. & A. Gray, Fl. N. Amer. 2:282. 1842. Berlandiera
pumila var. dealbata (Torr. & A. Gray) Trel., Rep. Ark. Geol. Survey 1888, 4:1
1891.

Berlandiera dealbata (Torr. & A. Gray) Small, Fl. S.E. U.S. 1246, 1340. 1903. Type:

NITED STATES. “Arkansas” [present day Oklahoma], [date], 7 Nuttall (HOLO-
TYPE: NY fiche!).

Berlandiera puma (Michx.) Nutt. var. scabrella Nesom & Turner, var.
nov. Type: U.S.A. Texas. Jasper Co.: roadside of Hwy 63, 0.8 mi E of jet with FM
1738, 1.0 mi E of crossing of Melholms Creek, 7.0 mi E of jct Hwys 290 and 96 in
Jasper; narrow strip of ie vegetation along side ee buffer to loblolly pine
plantation behind; dominant scrubby post oak, blackjack oak, black hickory, and
bluejack oak, with less common sweetgum, dogwood, sassafras, yaupon, buckeye,
and winged sumac; Berlandiera scattered but abundant along edge of woods for ca
50 yards on N side of hwy, in deep, loose sand; 16 May 1998, G.L. Nesom 98/1
(HOLOTYPE: TEX; isotypes: ASTC, BRIT, FSU, GH, IBE, LSU, MISSA, MO, NCU,

NLU, OKL, SHST, TAES, TAMU, UARK, US, USCH)
umilo vat. ilis sed differt vestimento hispidulo paginis superis foliaribus,

tr ‘chomar oie bas Blifeie erectis eae incrassitis.

Most of the variation that occurs among plants of Berlandiera pumila is
attributable to three patterns: 1) vestiture of the upper leaf surfaces—a relatively
abrupt transition in easternmost Texas from fine, thin-based, reclining or
appressed hairs to thick, basally erect hairs; 2) vestiture density of the stems
and lower leaf surfaces—interpopulational variation in central and east-central
Texas and the gradual reduction in density, with loosening of the matted
tomentum and shortening and thickening of the hairs, in populations from
west to east toward Louisiana; and 3) stem vestiture coloration—produc-
tion of reddish cross-wall pigments mostly in southeastern Texas and adja-
cent west-central Louisiana. Each of these patterns is discussed below in
more detail.

500 Sipa 18(2)

1) The morphological feature that shows the sharpest geographic transi-
tion within Berlandiera pumila is the nature of the vestiture on the upper
leaf surfaces: fine, silky, thin-based, basally reclining or appressed hairs (var.
pumila) vs. thicker-based and basally erect hairs (var. seabrella). It is prima-
rily on this basis that the taxonomic separation between the varieties of B.
pumila in the present interpretation can be made most consistently. Inter-
mediacy in hair morphology occurs in Cass, Harrison, Shelby, Polk, and
Tyler counties and Caddo and DeSoto parishes (and perhaps others). This
feature also was emphasized by Pinkava (1970) in his key to Texas Berlandiera
and was described by Torrey and Gray (1842, their “B. tomentosa y”).

2) Plants of Berlandiera pumila var. pumila with a dense vestiture of long,
thin, matted hairs show white stems and distinctly bicolored leaves (white
beneath, green above), the stem and lower leaf surfaces obscured by the
tomentum (the “white form” of var. pumila). Plants with trichomes short-
ened in length show a corresponding reduction in density of vestiture and
the green surfaces of the stems and leaves are visible through the hairs (the

“gray-green form’ of var. pumila, not mapped on Fig. 1, intermediate be-
tween the “white form” and “green form”). Shorter trichomes of the lower
leaf surfaces tend to be somewhat erect and separate rather than parallel to
the surface and closely intertwined. Only the white form of B. pumila oc-
curs in populations from the westernmost area of the range (Fig. 1). The
white form and gray-green form occur broadly over east Texas. In this area
are plants with dense white tomentum as well as plants with vestiture re-
duced by degrees in density, this feature tending to be relatively uniform
within a look population. A corresponding reduction in stem vestiture appears
to be the only mor phological difference correlated with the variation in
leaf vestiture. The “green form” of B. pumila occurs in easternmost Texas
counties and in Louisiana; in these plants the vestiture is further reduced,
with the lower leaf surfaces showing greenish or yellowish-green and hardly
different in color from the upper surface. In this area, the gray-green form
occurs infrequently and the white form is rare or absent. The reduction in
stem and abaxial leaf vestiture is characteristic of some populations of var.
pumila and all of var. scabrella. In the present attempt to characterize finer
patterns of infraspecific variation, we note the that the informally desig-
nated white, gray-green, and green forms are found within var. pumila oe

c

var. scabrella includes only the green form.

—

3) The stem vestiture of plants from a cluster of southeasternmost Texas
counties (Hardin, Polk, Jasper, Newton, Angelina, and Sabine cos.) and
adjacent Louisiana (Fig. 1) is reduced and the hairs distinctly colored. The
hairs are basally erect, though not thick and long like those of B. betonicifolia,
and the strongly colored cross-walls of all the hairs give a purplish or red-
dish-brown color to at least the upper stems. Such purplish stems are char-

NesoM AND Turner, The Berlandiera pumila complex 501

acteristic primarily of plants recognized here as var. scabrella, but intergra-
dation in this feature occurs with plants of var. pumila (the gray-green form)
from slightly further west. In these intergrades, hairs with colored cross-
walls are formed among the more abundant white hairs but are less notice-
able within the predominately grayish cauline vestiture.

The evolutionary origin of Berlandiera pumila var. scabrella, as suggested
by Pinkava (1977), may have been through hybridization between B. betonicifolia
and typical B. pumila. Both B. betonicifolia and B. pumila var. scabrella have
reduced vestiture of stems and leaves and neither produces white tomen-
tum; the upper leaf surfaces in both are somewhat hispid with thickened,
basally erect hairs. Stem hairs in both entities usually produce purplish cross-
walls. As noted above, however, such hybridization is not currently evident
and parallel evolution is an equally reasonable explanation of these mor-
phological similarities between B. betonicifolia and var. scabrella.

THE NATURE OF TYPICAL BERLANDIERA PUMILA

Berlandiera pumila in its typical form occurs in western Florida (where
the type collection was made by Michaux), Alabama, Georgia, South Carolina,
and North Carolina. The general distribution of the species was shown b
Pinkava (1967); its absence from Mississippi remains a reality (S. McDaniel,
pers. comm.). Various populations of B. pumila in this eastern segment of
its geographic range correspond to both the “white form” and “green form”
of the western segment of the species, as outlined in the comments above.
There is no variation within B. pumila of the eastern U.S. toward the mor-
phological features characteristic of var. scabrella.

KEY TO THE TAXA OF THE BERLANDIERA PUMILA COMPLEX

1. Stem hairs thick-based, erect, mostly 1.01. 5{-2 Ese long, commonly

distinctly flattened and vitreous; lower leaf

to deleate: the mid-cauline and upper usually gescile B. betonicifolia
1. Stem hairs soft and thin, densely matted and closely appressed or some-

times oriented in parallel, the length of the individual hairs difficul to
observe, sometimes more loosely aenee and less than 1 mm long, white;
lower leaf surfaces white to gray-green or greenish; leaves ovate to trian-
gular or lanceolate, the mid-cauline and upper usually on short (1-2
mm) petioles B. pumila (2)
2. Upper leaf surfaces softly pubescent with very thin, appressed or ba-
sally reclining hairs (sometimes basally erect in Cass, Harrison, and
Shelby cos., Caddo Parish), the leaves strongly to weakly or not at all
bicolored; stem vestiture relatively dense and matted or reduced in
density with the stem surface visible, the hairs evenly whitish or some
with purplish cross-walls near the base umila var. pumila
2. Upper leaf surfaces minutely hispidulous with relatively thicker-based
and basally erect hairs, the leaves not bicolored; stem surface visible,
the vestiture reduced in density, the hairs often reddish-brown to purple
because of colored cross-walls B. pumila var. scabrella

502 Stipa 18(2)
ACKNOWLEDGMENTS

Specimens have been examined from ASTC, BRIT, NLU, SBSC, SHST,
TAES, TAMU, and LL, TEX. The K holotype was loaned to Turner at LL, TEX.
Walter Holmes provided information on collections housed at BAYLU and
Sidney McDaniel verified the absence of Berlandtera pumila from Missis-
sippi. Thanks to Mark Bierner and Reviewer #2 for their helpful criticisms.

REFERENCES
Cronauist, A. 1980. Asteraceae. Vascular flora of the southeastern United States, Volume
I. Univ. of North Carolina Press, Chapel Hill.
GaNnbul, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. 4:1—

202.

Geiser, S.W. 1948. Naturalists of the Frontier (ed. 2). Southern Methodist University Press,
Dallas, Texas.

Gray, A. 1842. Berlandiera. In: Torrey, J. and A. Gray. Fl. N. Amer. 2:280-283.

Great PLAINS FLORA ASSOCIATION. 1986. Flora of the Great Plains. Univ. of Kansas Press,

awrence.

MacRoserts, D.T. 1989. A documented checklist and atlas of the vascular flora of Loui-
siana. Bull. Mus. Life Sci., No. 8:1-534; Appendix, bibliography, and addendum.

Orzett, §.L. and E.L. Bripces. 1987. Further additions and noteworthy collections in
the flora of Arkansas, with historical, ecological, and phytogeographical notes. Phytologia
64:81—-144.

Pinxava, D.J. 1967. Biosystematic study of Berlandiera (Compositae). Brittonia 19:285—

—

08.
. 1970. Berlandiera. Pp. 1623-1625. In: Correll, D.S. and M.C. Johnston. Manual
of the vascular flora of Texas. Texas Research Foundation, Renner, Texas.

SHINNERS, L.H. 1951. Notes on Texas Compositae—VIII. Field & Lab. 19:133-136.
SMALL, J.K. 1903. Flora of the southeastern United States. Published by the author, New

ork.

SMALL, J.K. 1933. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel
Hill.

SmitH, E.B. 1994. Keys to the flora of Arkansas. Univ. of Arkansas Press, Fayetteville.

Taytor, R.J. and C.E.S. Tayior. 1994. An annotated list of the ferns, fern allies, and flow-
ering plants of Caney (ed. 3). Biology Dept. Herbarium, Southeastern Oklahoma
State University, Dur

TORREY, - R. and A. Gray. 7 1842. A flora of North America, Vol. 2. part 2. Wiley and Putnam,
New

TURNER, B. L. and M.C. JOHNSTON. 1956. Chromosome numbers and geographic distri-
bution of Lindheimera, eC ae and Berlandiera (Compositae—Heliantheae—
Melampodinae). Southw. Nat. 1:125—132

NEW NAMES AND COMBINATIONS IN
NEOTROPICAL MYRSINACEAE

JOHN J. PIPOLY II

Botanical Research Institute of Texas
9 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
Jpipoly@brit.org; clusia@latinmail.com

JON M. RICKETSON

Missourt Botanical Garden
d - OX
St. Louis, MO 63166-0299, U.S.A.
jricketson @lehmann. mobot.org
ABSTRACT

Lundell (198 1a, 1986) published 68 new names and binations for binomials in Ardisia
and Myrsine. In both papers, he indicated that he was opposed to placement of the taxa
involved in those genera and that the combinations and new names were provided in an-
ticipation of future circumscriptions of the genera. We agree with Morales (1997) that in
doing so, Lundell inadvertently invalidated his new names and combinations, according
to the International Code of Botanical Nomenclature Article 34.1. To remedy the situation,
the following new proposed in Myrsine: M.. ee (Lundell) Ricketson
& Pipoly and M. juergensenti (Mez) Ricketson & Pipoly; the other seven Lundell combi-
nations are taxonomic synonyms and thus, not in need of validation. In addition to those
combinations validated by Morales (1997) in Ardisia, 27 new combinations are made, nine
new names are proposed, and for those taxa described by Lundell in the segregate genus
Icacorea Aublet but never transferred, five new combinations are proposed. The remaining
17 binomials are taxonomic synonyms and therefore, are not validated.

RESUMEN

Lundell (1981a, 1986) publicé 68 nombres y combinaciones nuevos para binomiales
en los géneros Ardisia y Myrsine. En ambos trabajos, el indicé que no estaba de acuerdo
con i Eble on re eomics de los taxa dentro de los géneros susodichos y que se estaba
llevan b nticipar circumscripciones futuras

Ps

que ee ser descritas ae los ees Esto: de acuerdo con Moret: (1997) quien
consideraba que Lundell, p
nuevos segun el Codigo nal de per Botanica (ICBN) OTe 34.1. Para
corregir esta situacién, las combinaciones nuevas se proponen dentro del género Myrsine:
M. calcarata (Lundell) Rickeson & Pipoly, y M. juergensenii (Lundell) Ricketson & Pipoly;
las otras siete combinaciones que hizo Lundell se consideran com taxonomicos

or lo tanto, no necesitan ser transferidos. Ademas de las combinaciones validadas por
Morales (1977) dentro de Ardisia, se proponen 27 combinaciones nuevas y nueve nombres

SipA 18(2): 503-517. 1998

504 SipA 18(2)

nuevos. Para los taxa descrito por Lundell dentro del géner d mo /cacorea Aublet

ion nunc “ tr ansferidos al gener oO Ardista, s se Ur ransfierren cinco et aciones nuevas. Los
17 y pot lo tanto, no necesitan ser tr ansferidos.
INTRODUCTION

Cyrus L. Lundell studied the neotropical members of the Myrsinaceae
for over 60 years, and was the principal monographer of the family since
the monograph of Mez (1902) in Engler’s Das Pflanzenreich. In addition
to treatments of the family for Flora of Guatemala (Lundell 1966) and Flora
of Panama ( (Lundell 1971), Lundell published a series of 26 papers in the
journals Wrightia and Phytologia dealing with the taxonomy of the family.
The senior author began collaboration with Lundell in 1981 while a graduate
student, and corresponded with him regarding generic delimitation for almost
six years.

Ardisia was described by Swartz (1788) in his Nova Genera et Species Plantarum,
and eventually conserved over the earlier /cacorea Aublet (1775). However,
Lundell perceived that Ardisia, as defined in the broad sense by Mez (1902),
was an artificial assemblage (paraphyletic in more modern terminology),
and therefore, segregated the following genera from it: Amatlania Lundell
(1982), Auriculardisia Lundell (1981c), Chontalesia Lundell (1982), Gentlea
Lundell (1964), Graphardisia (Mez) Lundell (198 1a), /barrea Lundell (198 1a),
Oerstedianthus Lundell (198 1b), Synardisia (Mez) Lundell (1963), Valerioanthus
Lundell (1982), Yunckeria Lundell (1964), and Zunilia Lundell (1981c).
While some of these segregate genera undoubtedly represent monophyl-
etic lineages, the question of their respective circumscriptions and the con-
sequent limits of the pantropical genus Ardisia, as well as the limits of all
the genera in the tribe Ardisieae, was often a source of confusion.

Simultaneously, circumscription of the genus Myrsine L. and its relationship
to the pantropical genus Rapanea Aublet, formed another source of taxo-
nomic debate. While Mez (1902) recognized Myrsine as a small genus of
paleotropical species, Hosaka (1940) demonstrated that the characters used
to separate the two were not constant, resulting in Walker's (1959) change
in taxonomic opinion. Lundell (1966, 1971) recognized Rapanea as dis-
tinct from Myrsine, but noted that Stearn (1969) elected to accept the more
inclusive concept of Myrsine to include Rapanea in his treatment of the Jamaican
species, a view increasingly adopted by modern workers (see Pipoly 1996
for a complete discussion).

During the early 1980s, the senior author was in close communication
with Lundell and frequently discussed the problem of generic delimitation
in the Myrsinaceae, particularly when viewed on a worldwide basis. The
problem was compounded when preliminary cladograms were tested, and
it appeared that several groups recognized by Mez were found to be paraphyletic
(Pipoly 1987). As Lundell carried out the mammoth job of describing the

PIPOLy AND RICKETSON, New names and combinations in Myrsinaceae 505

many neotropical novelties discovered in the late 1970's and early 1980’s,
it became necessary for him to determine limits of the genera, and he ap-
parently decided to provide names for the Ardisioid taxa both in the nar-
rowest and broadest concepts that had been circumscribed to that date.
Unfortunately, this philosophy led to the publication of two Papers in which
he essentially proposed new names and combinations in anticipation of future
circumscription, even though he was not convinced of the placement of
the taxa, thus invalidating all of them according to ICBN Article 34.1.

In preparation for our treatment of the Myrsinaceae for the Flora
Mesoamericana Project of the National Herbarium of Mexico (MEXU), Missouri
Botanical Garden (MQ), the British Museum (BM) and other collabora-
tors, we have published synopses for the genera Myrsine (Ricketson & Pipoly
1997a), Stylogyne (Ricketson & Pipoly 1997b), Gentlea (Ricketson & Pipoly
1997c), along with a revision of the genus Ardisia subgenus Graphardisia
Mez (Pipoly & Ricketson, 1998). The problems posed by Lundell’s (198 1b)
transfer of Mesoamerican Rapanea species to Myrsine were overlooked, but
are rectified below. As work continues on Ardisia in its broadest sense, it
becomes increasingly necessary to validate the invalid names and combi-
nations made by Lundell (1986) in transferring species described in the
various segregate genera of Ardisia to that genus, just as Morales (1997)
did for only a few from Costa Rica and Panama. The consequent combi-
nations, new names and updated synonymies for the taxa we recognize in
these two problematic taxonomic groups are also effected below.

NOTES ON MYRSINE

Lundell (198 1b) described the genera /barrea and Oerstedianthus as new,
raised Ardisia subgenus Graphardisia to generic status, and transferred the
Mesoamerican (and one Peruvian) species of Rapanea to Myrsine. How-
ever, in the introduction, Lundell stated “Nine species described in Rapanea
Aubl., all except one from Mexico and Central America, are transferred to
Myrsine L. to follow current practice, not out of any conviction as to the
justification for this disposition of the taxa!” By including that statement
in the introduction, he inadvertently invalidated the subject new combi-
nations because Article 34.1 of the ICBN unequivocally states, “A name is
not validly published (a) when it is not accepted by the author in the origi-
nal publication; (b) when it is merely proposed in anticipation of the fu-
ture acceptance of the group concerned, or of a particular circumscription,
position or rank of the group (so-called provisional names), except as pro-
vided for in Art. 59...” While there is no doubt that Lundell recognized
the entities involved as distinct taxa, we feel that he purposely effected the
transfers as a result of discussions with the senior author and in anticipa-
tion of his (and others’) future work on the group as the more broadly cir-

506 Sipa 18(2)

cumscribed Myrszne rather than setting it apart as Rapanea. Therefore, among
the nine taxa he transferred, all are invalid. However, Ricketson and Pipoly
(1997a) recognized only two of these species, Myrsine calcarata and M.
guergensenii. The other six species, Rapanea pittieri, R. peruviana, R. rufa,
R. panamensis, R. allenii, R. mexicana and R. reflexiflora, were synonymized,
and thus the combinations made by Lundell are nomenclatural synonyms
of them, with no need for validation. The new combinations and corrected
synonymies for those species are presented below. For complete synony-
mies and citation of basionyms of recognized taxa, see Pipoly and Ricketson

(1997a).

Myrsine calcarata (Lundell) Ricketson & Pipoly, comb. nov. Rapanea calcarata
ee ee 295.1976. se iets il (Lundell) Lundell, Phytologia 48:142.
198 omen invalidum. TYPE: COS ‘A. ALAJUELA: S side of Volcan Pods,
over oe on road leading past — station, without elev., 24 Mar 1972, D
Stone 3141 (HOLOTYPE: LL-TEX).

Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult. subsp. coriacea, Syst.
Veg. 4:511. 1819.

ie fel Lundell, Wrightia 5:298. 1976. melas rufa (Lundell) Lundell, Phytologia
81,nomen invalidum. Type: COSTA RICA. SAN JOSE: Cordillera de Talamanca,
ee ae se of the Chirrip6é Massif, 2,700—3,000 m, 6 Apr 1969, G. Davidse & R.
Pohl 1635 (HoLoryre; LL-TEX; tsorype: MO)
Myrsine dependens (Ruiz & Pav.) Spreng., Syst. Veg. 1:664. 1825.

pittiert Mez in Engl., Pflanzenr. 7 236(Heft 9):378. 1902. Myrsine pittieri
ez) Lundell, Phytologia 48: 142. 1981, nomen invalidum. Type: COSTA RIC
eee Volcan Iraza, 3,000 m, H. ae ae (HOLOTYPE: B-destr. 1943; LECTO-
TYPE: by Pipoly 1992, BR; IsoLecroryPe:

Rapanea peruviana Lundell, Wrightia 6:117. 1980. Myrsine pernviana (Lundell) iets

Phytologia 48:142. 1981, nomen invalidum. Type: PERU. Cuzco: Tres Cruces, up-
per edge of fe gas Nacional de Manu, 1-13 km NW of Paucartambo-Pilcopata ead.
3,300—3,50( 29 Jun 1978, A. Ps al. 23478 (HOLOTYPE: LL-TEX; IsoryPEs:
MO, ty

Myrsine juergensenti (Mez) Ricketson & Pipoly, comb. nov. Rapanea juergensenii

Mez in Engl. erent IV. 236(Heft 9):388. 1902. Myrsine juergensenti (Mez) Lundell,
Phycologia 48:1 1981, nomen invalidum. TYPE: MEX ae AXACA: without ae

without elev., without date, H. Galeotti 530 (LECTOTYPE: US, by Ricketson & Pipoly,
1997a; ISOLECTOTYPES: G, NY).

z=

Rapanea allenii Lundell, Wrightia 4:168. 1971. Myrsine allenii (Lundell) Lundell fees
48:142. 1981, nomen invalidum. Type: PANAMA. Cuiriqut: N forested fac rro
Copete, an E spur of Chiriqui Re 2,438—-2,591 m, 29 Jul 1947, P. An 7
(HOLOTYPE: US; tsoTypes: G O).

Rapanea mexicana Lundell, eae 5:296. 1976. Myrsine mexicana (Lundell) Lundell,
Phytologia 48:142. 1981, nomen invalidum. Type: MEXICO. Cuiapas: Pico de Loro,
near Escuintla, without elev., 25 Jun 1941, E. Matuda 4228 (nototyPE: LL-TEX;
IsOTYPES: A, F, , US).

Rapanea reflexiflora Lundell, Wrightia 5:297. 1976. Myrsine reflexiflora (Lundell) Lundell,

PIPOLY AND RICKETSON, New names and combinations in Myrsinaceae 507
Phytologia 48:142. 1981, nomen invalidum. TYPE: PANAMA. COCLE: Woods ad-
jacent to chicken farm, La Mesa, above El Valle, without elev., 3 Jan 1974, J, Dwyer
11885 (HOLOTYPE: LL-TEX; ISOTYPES: BRIT, KF MO, PMA).

nine ellucidopunctata ae De Meddel. Dansk Naturhist. Foren.

Kjobenhavn 1861: 133. 1
Kapanea tera I sa Wrightia 4:169. 1971. saad ‘panamensis (Lundell) Lundell,
Phytologia 48:142. 1981, nomen invalidum. Tyre AMA. PANAMA: San José
Island, Berlas ee Gulf of Panama, ca. 55 ral SSE of Balboa, main road W
of Matta Puerco, 9 Nov 1944, /. Johnston 441 (HOLOTYPE: LL-TEX; isorypes: GH,
MO, US).

NOTES ON ARDISIA

The situation regarding generic delimitation among the taxa compris-
ing Ardisia in its broadest sense, and that of the segregate taxa proposed or
recognized by Lundell, was very similar to that described above for Myrsine.
Work underway among these taxa for our studies in Mesoamerica has caused
us to reexamine the nomenclatural legalities regarding the new combina-
tions and names proposed by Lundell (1986), particularly in light of the
recent new names and combinations made by Morales (1997), in his paper
see a treatment of the Myrsinaceae for the A Manual to the Plants
of Costa Ri

fandall (1986) effected 59 nomenclatural changes, comprising 23 new
names and 36 new combinations for taxa previously described in genera
segregated from Ardisia. However, in the introduction, he stated, “Just as
the very distinct genus /cacorea Aubl. has been ridicuously submerged as a
subgenus of Ardisia Sw. for generations, we can also assume that the vagar-
ies of taxonomists will result in the reduction of such recently described
genera as Auriculardisia Lundell, Gentlea Lundell, Graphardisia Lundell,
Oerstedianthus Lundell, [barrea Lundell, Amatlantia Lundell, and Zunilia
Lundell to subgenera of Ardisia. The transfer of the following species is made
to that genus to obviate the incentive of workers to undertake such unnec-
essary reductions to subgenera.” While the wording of this introduction is
not as clear a violation of the rules of Article 34.1 of the ICBN as that of
the Myrsine paper, we feel that the intention was to specifically provide names
“proposed in anticipation of the future acceptance of the group concerned,
or of a particular circumscription, position or rank” and are therefore, in-
valid. This agrees with the conclusion reached by Morales (1997) in his
validation for six of the subject taxa. In addition, while implied and not
specifically stated, we feel that Lundell accepted the taxa, but not the names
(i.e., the placement) he published in the work.

Our work thus far has shown that among the segregate genera recog-
nized by Lundell, Gentlea, Ctenardisia Ducke (= Yunckeria Lundell), and

Synardisia will all be recognized with certainty, while the remaining groups

508 Sipa 18(2)

will not. Because we recognize Gentlea as a genus distinct from Ardisia, no
validation of the invalidly published Lundell new names and combinations
for Gentlea species as members of Ardisia is necessary, but we provide the
following corrections to synonymies listed in our previous work, to include
the invalid names (Ricketson & Pipoly 1997c). For complete synonymies,
see Ricketson and Pipoly (1997c).

Gentlea austinsmithii (Lundell) Lundell, Wrightia 4:68. 1968. Gentlea molinae

Lundell, Wrightia 4:150. 1970. Ardisia molinae (Lundell) Lundell, Phytologia 61:65.

1986, nomen invalidum. TYPE: HONDURAS. LA PAZ: Montafia Verde on Cordil-

lera Guajiquiro, 1,900 m, 23 Mar 1969 (fr), A. Molina R. & A. Molina 24389 (HO
LOTYPE: LL-TEX; [SOTYPES: EK LL-TEX, NY).

Gentlea parviflora Lundell, Wrightia 5:89. 1975, non Ardisia parviflora Talbot, Syst. List
‘Trees, Bombay ed. 2, 204. 1902. Ardisia parvissima Lundell, Phytologia 61:66. 1986,
nomen novum invalidum Tyre: GUATEMALA. Baja Verapaz: Union Barrios, in high
forest, on top of hill, E of km 154, 11 oy 1975 (fl), C. Lundell & E. Contreras 19212
(HOLOTYPE: LL-TEX; isotypes: CAS, ‘

Gentlea pase Lundell, ace 6:96. 1979, Non Ardisia eeu Vent.,
Choix PL., 5. 1803, non Ardisia crenulata C. Lodd., Bot. Cab.
nom. illegit. Ardisia contrerasti Lundell, Phytologia 61:63.1986, nomen novum invalidum.
TYPE: GUATEMALA. BAJA VERAPAZ: Nifio Perdido, bordering Rio San José, 8 km
N, in high forest, on top of hill, 27 May 1977 (Al, my C. Lundell & E. Contreras 21004
(HOLOTYPE: LL-TEX; ISOTYPES CAS, LL-TEX 2-SHEETS)

2

—

Gentlea lancifolia Lundell, Phytologia 58:273. 1985. Non Ardisia lancifolia
Merr., Philipp. J. Sci. 13:113. 1918. Ardisia intibucana Lundell, Phytologia 61:65
1986, nomen novum invalidum. TYPE: HONDURAS. INTIBUCA: Between Calaveras
and El Duraznillo, Cordillera Opalaca, 1,800 m, 12 Mar 1970 (fr), A. Molina R. &
A, Molina 25547 (HOLOTYPE: F; ISOTYPE US).

Gentlea micranthera (Donn. Sm.) Lundell, Wrightia 4:69. 1968.

Gentlea cuneifolia Lundell, Wrightia 7:245. 1983. Ardisia cuneifolia (Lundell) Lundell,
Phytologia 61:63. 1986, nomen invalidum. Type: MEXICO. Veracruz: Municipio
de San Andrés Tuxtla, ca. del aguaje en el lado S del Volcan San Martin Tuxtla, 1,250
m, 15 Feb 1972 (fr), /. Beaman & C. Alvarez del Castillo 5751 (HoLotypE: LL-TEX;
IsOTYPES: F, MEXU).

Gentlea auriculata Lundell, Phytologia 58:273. 1985, non Ardis Donn. Sm.,
Bot. Gaz. 24:395. 1897. Ardisia parviauriculata Lundell, rea 61:66. 1986,
nomen novum pvelcun. 7 GUATEMALA. Zacapa: Sierra de Las Minas, middle
and upper S-facing slopes of Volcan Gemelos, 2,100-3,200 m, 26 Jan 1942 (fl, fr), 7.
Steyermark 43277 (Hi HOLOTYPE F; tsorypes: LL-” ;

Gentlea maculata Lundell, Phytologia 58: 374, 1985. Non Ardisia maculata .
Poit. ex A. DC., Trans. Linn. Soc., London 17:116. 1834. Ardisia morazanensis Lundell,
Phy tologia 61: 65. 1986, nomen novum invalidum. TYPE: eclrr ae FRANCISCO
MORAZAN: On mountain La Tigra, SW of San Juancito, 1,800—2,100 m, 2 Feb 1966

fl), A. Molina R., L. Williams, W. Burger & B. Wallenta 16972 (HOLOTYPE: F; ISOTYPES:

, US).

Ter
a

Gentlea standleyi Lu ndell, Wrigh tla 4:69. 1968. Ardisia standley: (Lundell) Lundell,
Phytologia 61:67. 1986, nomen invalidum. TYPE: COSTA RICA, HEREDIA: Cerros

PIPOLY AND RICKETSON, New names and combinations in Myrsinaceae 509

de Zurqui, NE of San Isidro, 2,000—2,400 m (fr), 3 Mar 1926, P Standley & J. Valerio
DOGIS (HOLOTYPE US AsOTYPE: LiL- TEX),

Gentlea venosissima (Ruiz & Pavén) Lundell, Wrightia 3:103. 1964.

Gentlea tenuis Lundell, Wrightia 7:24. 1981, non Ardisia tenuis Lundell, Wrightia 4:149.
Ardisia jitotolana Lundell, Phytologia 61:65. 1986, nomen novum invalidum
Type: MEXICO. Cuiapas: Municipio de Rayén, in the Selva Negra 10 km above Rayén
Meycalapa along road to Jitotol, 1,700 m, 12 Dec 1971 (fl), D. Breedlove 23144 (Ho-
LOTYPE: F; isoTypes: DS, LL-TEX, MO, NY
While we agree with Morales (1997) in his view that the new names and
combinations published by Lundell (1986) are invalid, Morales did not
make direct reference to the invalid names in the synonymy listed with new
combinations. Therefore we provide them below in the interest of provid-
ing complete synonymy for those species.

Ardisia apodophylla J.E Morales, Lees 83: 111. 1997 [1998]. kien
sessilifolia Lundell, Wrightia 7:272. 1984, n Engler, Pfan
IV. 236(Heft 9):125. 1902. pce Lundell, Phytologia 61:68. 1986, nomen
novum invalidum. Type: COSTA RICA. ALaJUELa: Cordillera Central near San Juan
de Laja about 15 km N of Zarcero, 1,350 m, 7 Feb 1965 (fl, fr), L. Williams, A.
Molina R., T. Williams & D. Gibson 28998 (HOLOTYPE: F).

Ardisia bekomiensis ee E Morales, Phytologia 83:111. 1997 [1998].
komiensis Lundell, Phytologia 56:414. 1984. Ardisia bekomiensis (Lundell)
sali Seer 62. 1986, nomen invalidum. Type: COSTA RICA. PUNTARENAS—
Limon: border, Cordillera de Talamanca, Cerro Bekom, 11 airline km SSW of the
peak of Cerro Kamuk, 9° 10' 30" N, 83° 03' 45" W, 2,6002,750 m, 21 Mar 1984
(fl), G. Davidse, G. Herrera Ch. & R. Warner 25710 (HoLotyre: LL-TEX; IsoryPEs:

INB, MO).

Ardisia eurubiginosa (Lundell) J.E Morales, Phytologia 83:111. 1997 [1998].
Auriculardisia eurubiginosa Lundell, Phytologia 56:413. 1984. Ardisia eurubiginosa
(Lundell) Lundell, Phytologia 61:63. 1986, nomen invalidum. Type: PANAMA. Bocas
DE Toro: Cordillera de Talamanca, 2—5 airline km NW of the Peak of Cerro Echandi
on the Costa Rican-Panamanian international border, 2,600—2,850 m, 1—9 Mar 1984
(fl, fr), G. Davidse, L. Gémez, G. Herrera, C. Chacén, I. Chacon & A. Chacén 25486
(HOLOTYPE: LL-TEX; isoryres: INB, MO, NY).

Ardisia quadrata (Lundell) J.F. Morales, eee 83: il 1 L. 1997 [1998].
Auriculardisia quadrata Lundell, Phytologia 56:413 Adrata (Lundell)
Lundell, Phytologia 61:66. 1986, nomen invalidum. ae Coun RICA. PUNTARENAS:
Foothills of the Cordillera de Talamanca, around Tres Colinas, 9° 07' N, 83° 04' W,
1,800-1,850 m, 20 Mar 1984 (fl) G. Davidse, G. Herrera Ch. & R. Warner 25645
(HOLOTYPE: LL-TEX; 1sotyres: INB, MO)

Ardisia sordida (Lundell) J.Z Morales, Phytologia 83:111. 1997 [1998].
Auriculardisia sordida Lundell, Wrightia 7:272. 1984. Ardisia sordida (Lundell) Lundell,
Phytologia 61:66. 1986, nomen invalidum. Type: COSTA RICA. ALajuELa: Finca
Los Ensayos, ca. 11 mi. NW of Zarcero, ca. 850 m, 15 Aug 1977 (fl), 7) Croat 43538
(HOLOTYPE: LL-TEX; 1soTyres: CR, MO)

Ardisia squamata (Lundell) J.K Morales, Phytologia 83:111. 1997 [1998].
Auriculardisia squamata Lundell, Phytologia 56:19. 1984. Ardisia squamata (Lundell)

510 Sipa 18(2)
Lundell, Phytologia 61:67. 1986, nomen invalidum. Type: COSTA RICA. HEREDIA:
Original forest near the Rio Puerto — ca. 2 km upstream from the confluence
with Rio Sarapiqui, formerly “Finca La Selva” of L.R. Holdridge, 10° 26' N, 84° 0°
Ww, 100 1 m, 1417 oe 1968 8 (fl), W. ee e® R. Stolze 5853 (HOLOTYPE: F; ISOTYPES:
CR, EK TEX, M

Regarding those taxa originally placed by Lundell (198 1a) in his concept

of the segregate genus Graphardisia, the following binomials invalidly published

by Lundell (1986) as new combinations and names: Ardisia coibana (Lundell)

Lundell, Ardisia ebingeri Lundell, Ardisia esquipulasana Lundell, Ardisia neohyalina

Lundell, Ardisia obtusata (Lundell) Lundell, Ardisia riomonteana Lundell,

Ardisia seranoana (Lundell) Lundell and Ardisia ustupoana (Lundell) Lundell,

are considered taxonomic synonyms (Pipoly & Ricketson 1998) and thus,

need no validation.

Lundell (1986) also described two new species of Ardisia, A. hirsutissima
Lundell, and A. tarariae Lundell. He aligned Ardisia hirsutissima with A.
nevermannii and A. ursina, two species he had previously transferred (Lundell
1982) to his segregate genus Valerioanthus, and subsequently transferred
them once more, to his segregate genus Auriculardisia (Lundell 1985). Curiously,
Lundell (1987) transferred Ardisia hirsutissima to Valerioanthus rather than
to Auriculardisia, as he had previously done with the other members of
Valerioanthus and which, at that point, had its type already transferred. It
is unclear whether these nomenclatural transactions ene a mere dapsus
calimae, or a purposeful taxonomic decision. Lundell (1987) validly trans-
ferred Ardisia tarariae Lundell to Auriculardisia. However, our studies have
shown that this taxon ts also best retained in Ardisia. In order to provide
complete synonymies for these apparent taxonomic nomads, we provide
the following nomenclator.

Ardisia hirsutissima Lundell, Phytol eel 61:64. 1986. Valerioanthus hirsutissima
(Lundell) Lundell, Phycologia 63:78. 1987. Type: PANAMA. Coc Le: Continenta
Divide above El Copé, 8° 38’ N, 80° 39' W, 650750 m, 27 Nov 1985 (ft 1), G. de
Nevers, A. Henderson, H. Herrera, G. McPherson & L. Brako 6408 (HOLOTYPE: LL-
TEX; tsorypes: BM, CAS, CR, MEXU, MO, NY, PMA).

Ardisia nevermannii Standl., J. Wash. Acad. Sci. 17:524. 1927. Valerioanthus
nevermannu (Standl.) Lundell, Wrightia 7:50. 1982. Auriculardisia nevermannii (Standl.)
Lundell, Phytologia 57:450. 1985. Type: COSTA RICA, LIMOn: Finca Montecristo,
on the Rfo Reventazén, below El Cairo, ca. 25 m, 18-19 Feb 1926 (fl), P Standley
& J. Valerio 48603 (HoLotyre: US)

Ardisia ursina Lundell, Wrightia 6:97. 1979. Valerioanthus ursinus (Lundell) Lundell,
Davee 7:50. 1982. Auriculardisia ursinus (Lundell) Lundell, Phytologia 57:450.
1985. Type: PANAMA. Panama: El Llano-Carti Road, 10 km from Inter-American
Hwy., without elev., 5 Oct 1974 (fl), S. Mort & J. Kallunki 2314 (HoLotYPE: MO),

Ardisia tartariae Lundell, Phytologia 61:67. 1986. Auriculardisia tartariae (Lundell)
Lundell, Phytologia 63:75. 1987. Type: COSTA RICA. Limon: Cordillera de Talamanca,

i)

PirpOLy AND RICKETSON, New names and combinations in Myrsinaceae Sd

Atlantic slope, Cerros Tararia (locally known as Tres Picos), 2,400-2,600 m, 10 Sep
1984 (fl), G. Davidse, G. Herrera Ch. & M. Grayum 28882 (HOLOTYPE: LL-TEX;
IsoTyPE: MO).

For the remaining 27 binomials invalidly published by Lundell (1986), we
provide the following new names and combinations.

Ardisia alajuelae (Lundell) Pipoly & Ricketson, comb. nov. Icacorea alajuelae
Lundell, Phytologia 56:141. 1984. Ardisia alajuelae (Lundell) Lundell, Phytologia
61:62. 1986, nomen invalidum. Tyre: COSTA RICA. ALaAJUELA: San Pedro de San
Ramen. without elev., 12 Mar 1932 (fl), A. Brenes 15091 oe PE: ne

Ardisia albipedicellata Pipoly & Ricketson, nom. nov. eacorea parvifolia Lundell,
Phytologia 57: ae mire non sia aia Humb. ex Willd, in Rosin: & Schult,
Syst. Veg. 4:804. 1 Ardisia albiped: indell, Phytologia 61:62. 1986, nomen
novum ae i pe: COSTA RICA. PUNTARENAS: Just E of Monteverde on the
Pacific watershed, 10° 18' N, 84° 48' W, 1,3001,450 m, 29 Oct2 Nov 1975 (fl), W
Burger & R. Baker 9628 (HOLOTYPE: F).

Ardisia albisepala (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
albisepala Lundell, Wrightia 7:266. 1984. Ardisia albisepala (Lundell) Lundell, Phytologia
61:62. 1986, nomen invalidum Tyrer: PANAMA. Veracuas: Trail on ridge to sum-
mit of Cerro Tute, Cordillera de Tute, 1 km past Escuela Agricola Altos de Piedras,
W of Santa Fe, 8° 36' N, 81° 06' W, 9501,250 m, 15 Dec 1981 (fl), S. Knapp y K.
Sytsma 2548 (HOLOTYPE: LL-TEX; isotypes: MO, NY)

Ardisia azaharensis Pipoly & Ricketson, nom. nov. Auriculardisia microcalyx
Ey angel Wrightia 7:270. 1984, non Ardisia microcalyx Lundell, Wrightia 4:46. 1968.

wzaharensis Lundell, Phytologia 61:62. 1986, nomen invalidum. ‘Type: COSTA

RICA. ALajueLa: 15 km NW of San Ramén by air, Cerro Azahar, headwaters of Rio

San Pedro, by road, 9 km NW of San Ramén to Piedades Norte, then 3 more km

W to La Paz, 1.7 km to cluster of houses, then left again on jeep road 45 km to

top of ridge, 10° 09' 30" N, 84° 3435' W, 1,4001,500 m, 14 May 1983 (fr), &.

Liesner, E. Judziewicz, J. Gémez-Laurito, B. Pérez G. & A. Carvajal 15575 (HOLo-
TypE: LL-TEX; isotypes: k MQ).

an patent Pipoly & Ricketson, nom. nov. Auriculardisia par ee Lundell,
ightia 7:271. 1984, non Ardisia parviflora Talbot, Syst. list Trees, Bor
Ns 1902. Ardisia bristanii Lundell, Phytologia 61:62. 1986, nomen novum ‘invalidum
Type: PANAMA. Darien: Cerro Pirre, without elev., 4 Aug 1967 (fl), MV. Bristan
1236 (HOLOTYPE: US; IsoTyPe: MO).
Ardisia chiriquiana (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
ae Welaon a coe Wrightia 7:267. 1984. Ardisia chiriquiana (Lundell) Lundell,
g 1986, nomen eee Type: PANAMA. CHiriQut: Trail up Cerro
Pie Macho, 8° 50' N, 82° 25' W, 1,5001,900 m, 7 Jan 1938 (fl), B. Stein, B. Schmatlzel
e D. Roubik 1223 oer: L L-TEX; isotype: MQ).

Ardisia sane Pipoly & Ricketson, nom. nov. Auriculardisia wits Lundell,
Wri a 7:271. 1984, non Ardisia rosetflora Pitard, Fl. Indo-Chine, 3:866. 1930
Ardisia dhs oe Phytologia 61:63. 1986, nomen novum invalidum. Type:
PANA E: Trail from Continental Divide near the sawmill above El] Copé
to Rio Blanco del Norte, 8° 40' N, 80° 36' W, 350700 m, 20 Feb 1982 (fl), S. Knapp,
J. Mallet & R. Dressler 3646 (HOLOTYPE: MO)

512 Sipa 18(2)

Ardisia dryeri Pipoly & Ricketson, nom. nov. Auriculardisia micrantha Lundell,
Wrightia 7:269. 1984, non Ardisia micrantha Kunth, Nov. Gen. Sp. 3:246. 1818
[1819], non Ardisia micrantha Donn. Sm., Bot. Gaz. 14:26, 1889, nom. illeg. Ardisia
dryeri Lundell, Phytologia 61:63. 1986, nomen invalidum. Type: COSTA RICA.
ALAJUELA or PUNTARENAS: Reserva de Monteverde, Cordillera de Tilaran, al lado del
camino por la ventana, Div. Continental, 1,560—-1,580 m, 15 Aug 1976 (fl), Vv Dryer
577 (HOLOTYPE: MQ; ISOTYPE: F).

a

Ardisia eciliata Crane Pipoly & Ricketson, comb. nov. Zunilia eciliata Sean
Phytologia 58:490. 1985. Ardisia eciliata (Lundell) Lundell, s yroleee a 61:63. 19
nomen invalidum. - ype: MEXICO, Cuiapas: Municipio de € itla de Espinosa
Cerro del Ocote, 30 km NW of Ocozocoautla, 1,500 m, 14 Oct 1972 (fl), D. Breedlove
28973 (HOLOTYPE: LL-TEX; isorypes: CAS-DS, MEXU, MQ).

Ardisia arco Pipoly & Ricketson, nom. nov. asia ee Lundell,
Phytologia 56:19. 1984, non Ardisia elliptica Thunb., 71-8119, 179
Ardisia Mipofoli Lundell, Phytologia 61:63. 1986, nomen novum eae TYPE:
AXACA: Municipio de Comaltepec, km 149 carr. Tuxtepec, Sierra Juarez,
Puerto E ligio, 800 m, 17 Jun 1966 (fr), G. Martinez C. 884 (HOLOTYPE: XAL; ISOTYPES:
MEXU, MO).

In choosing the epithet ‘ellipticifolia’ over ‘elliptifolia’ used by Lundell, we
believe we are correcting a spelling error rather than changing the stem.

Ardisia eucuneata (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
eucuneata Lundell, Phytologia 57:449. 1985. Ardisia eucuneata (Lundell) Lundell,
Phytologia 61:63. 1987, nomen invalidum. Type: PANAMA. SAN BLas: Nusagandi,
trail from camp NW to a Quebrada, 9° 19’ N, 78° 15’ W, 300 m, 31 a 1984 (fl),
G. de Nevers & C. de Leon 3598 (HOLOTYPE: LL-TEX; 1soryre: MO)

——

Ardisia — Pipoly & Ricketson, nom. nov. Zumilia Hae = ll, Pe
1985, non Ardisia purpusti Brandeg., Univ. Calif. Publ. Bot. 6:1
a Jad Lundell, Phytologia 61:64. 1986, nomen novum inva ca 7 YPE:
MEXICO. Cuiapas: Mountains near Fenia, without elev., May 1925 (fr), C. Purpus
100 (HOLOTYPE: US)

Ardisia guanacastensis (Lundell) Pipoly = Nes comb. nov. /cacorea

guanacastensis Lundell, Phytologia 56:415. 4. Ardisia guanacastensis (Lundell)
Lundell, Phytologia 61:64, 1986, nomen nee Type: COSTA RICA, GUANACASTE:
Parque Nacional Rincon de la Vieja, the S$ E slopes of Volcan Santa Maria, aren

Estacion Hacienda Santa Maria, 10° 47' N, 85° 18' W, 9001,200 m, 2728 Jan 1°
(fl), G. Davidse, L. Gomez, M. Sousa, C. Humphries, N. Garwood, R. nee em
M. Gibby 23307 (HoLotYPE: LL-TEX; tsorypes: MO, NY).

Ardisia guinealensis (Lundell) Pipoly & Ricketson, comb. nov. Jeacorea guinealensis
Lundell, Phytologia 56:416. 1984. Ardisia guinealensis (Lundell) Lundell, Phytologia
61:64. 1986, nomen invalidum. Type: COSTA RICA. PUNTARENAS: Foothills of the
Cordillera de Talamanca, vicinity of Helechales, along the Rio Guineal, 9° 04' 30"

, 83° 05' W, 1,1001,200 m, 29 Mar 1984 (fr), G. Davidse e G. Herrera Ch. 26215
(HOLOTYPE: LL-TEX; tsoryre: MO)

Ardisia heterotricha (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
heterotricha Lundell, Wrightia 7:268. 1984. Ardisia heterotricha (Lundell) Lundell,
Phytologia 61:64. 1986, nomen invalidum. Type: PANAMA, PANAMA: On road near

PipOLy AND RICKETSON, New names and combinations in Myrsinaceae 513

slopes of Cerro Jefe, 2,400 ft [732 m
Putz 3417 (HOLOTYPE: MO).

Ardisia hornitoana Pipoly & Ricketson, nom. nov. /cacorea reflexa Lundell, Phytologia
56:20. 1984, non Ardisia reflexa Wall., Cat. No. 2282. 1830. Ardisia hornitoana Lundell,
Phytologia 61:64. 1986, nomen novum invalidum. Type: PANAMA. Cuiriqut: Path
from Linares farm ca. 1,400 m to top of Cerro Hornito, 1,750 m, 27 Dec 1977 (fl),
J. Folsom, R. Dressler & R. Channell 7257 (HoLotyre: LL-TEX; tsorype: MO).

Ardisia hugonensis (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
Augonensis Lundell, Wrightia 7:268. 1984, Ardisia hugonensis (Lundell) Lundell, Phytologia
61:64. 1986, nomen invalidum. Tyre: COLOMBIA. Cuoco: Municipio de Quibdé
Corregimiento de Guayabal, Rio Hugén, without elev., 12 Sep 1976 (fr), E. Forera
& R. Jaramillo 2812 (HoLoryrr: NY; isotype: MO)

Ardisia ixcanensis (Lundell) Pipoly & Ricketson, comb. nov. /cacorea ixcanensis

ell, Phytologia 58:489. 1985. Ardisia ixcanensis (Lundell) Lundell, Phytologia
61:65. 1986, nomen invalidum. Type: MEXICO. Cuiapas: Municipio ee
12 km from Laguna Tsiskaw on road to Ixcan, 1,250 m, 19 ‘Oa 1974 (fl), D
Breedlove 38965 (HOLOTYPE: CAS-DS).

pipisl jaliscensis (Lundell) Pipoly & Ricketson, comb. nov. /eacorea jaliscensis
indell, Phytologia 53:412. 1983. Ardisia jaliscensis (Lundell) I ae ee ogia
61: 65. 1986, nomen pivaiiduens Type: MEXICO. Jasco: Along road to Jirosto,
ca. 10 km WNW of Purificacién, 19° 44' N, 104° 42' W, 400 m, 11 are 1979 (fr),
H. lltis & M. Nee 1437 (HoLoTYPE: LL-TEX; tsoryre: US).

Ardisia knappii (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia knappii
Lundell, Phytologia 55:235. 1984. Ardisia knappii (Lundell) Lundell, Phytologia
61:65. 1986, nomen invalidum. Type: PANAMA. SAN Bias: 23—29 km from Pan-
American Highway on El Llano-Carti Road, 9° 22' N, 78° 69' W, 300400 m, 28
Oct 1981 (fl), S. Knapp 1843 (HOLOTYPE: LL-TEX; isotype: MO).

Ardisia latisepala (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
latisepala Lundell, Wrightia 7:269. 1984, non Icacorea latisepala Lundell, Phytologia
56:417. 1984. Ardisia latisepala (Lundell) Lundell, Phytologia 61:65. 1986, nomen
invalidum. Type: COSTA RICA. PUNTARENAS: 6 km S of San Vito de Java, around
Wilson’s finca, ca. 4,000 ft [1,219 m], 19 Aug 1967 (fl), P Raven 21653A (HOLO-
TYPE: F; isorype: MO).

ra

, 20 Jan 1980 (fr), 7) Antonio, H. Moore & E

Ardisia leptopoda (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
leptopoda Lundell, Phytologia 57:450. 1985. Ardisia leptopoda (Lundell) Lundell,
Phytologia 61:65. 1986, nomen invalidum. Tyre: PANAMA. Cuiriqut: E of Guadeloupe
along the Rio Chiriqui Viejo, ca. 2 mi NE of Cerro Punta, ridge of Cerro Respinga,
ca. 7,000 ft [2,134 m], 13 Jan 1971 (fr), R. Wilbur, J. Teeri & R. Foster 13111 (Ho-
LOTYPE: F),

Ardisia monteverdeana (Lundell) Pipoly & Ricketson, comb. nov. Leacorea
monteverdeana Lundell, Phytologia 57:451. 1985. Ardisia monteverdeana (Lundell)
Lundell, Phytologia 61:65. 1986, nomen invalidum. Type: COSTA RICA. PUNTARENAS:
Monteverde, Laguna-Hollander’s & Hoge’s, 1,320 m, 10 Dec 1979 (fl), S. Koptur
SK-251 (HOLOTYPE: LL-TEX; tsoType: MO).

Ardisia nebulosa (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia
nebulosa Lundell, Wrightia 7:270. 1984. Ardisia nebulosa (Lundell) Lundell, Phytologia
61:65. 1986, nomen invalidum. Type: PANAMA. PANAMA: Cerro Jefe, 850-900 m,
29 Oct 1980 (fr), K. Sytsma 1980 (HOLOTYPE: LL-TEX; isotypes: BM, M

514 Stipa 18(2)

Ardisia neomirandae me & Ricketson, nom. nov. Zunilia as Lundell,
Phytologia 58:491. 1985, non Ardisia mirandae Merr., Philipp. J. S 54.1
Ardisia neomirandae Lundell, Phytologia 61:66. 1986, nomen novum invalidum.
Type: MEXICO. Cuiaras: Finca Prusia, por camino a Mapastepec hasta—ca. Triunfo

(S. Jaltenango), without elev., 24 Feb 1951 (fl), F Miranda 7004 (HoLorype: MEXU).

Ardisia oaxacana (Lundell) Pipoly & Ricketson, comb. nov. eacorea oaxacana
Lundell, Wrightia 7:48. 1982. Ardisia oaxacana (Lundell) Lundell, Phytologia 61:66.
1986, nomen invalidum. Type: MEXICO. Oaxaca: ag Pluma Hidalgo, carretera
Pochutla, Oaxaca, 1 km al N de Chacalapa, 13 km al N de Pochutla, 225 m, 13
Nov 1979 (fl), S. P. Fryxell y 1! Wendt 79512 (HoLotyre: LL-TEX; Isoryees: F,
MO, NY).

—

Ardisia parvidenticulata Pipoly & Ricketson, nom. nov. Icacorea denticulata
Lundell, Phytologia 56:415. 1984, non Ardisia denticulata Blume, Bijdr. 13: 691.
1820. oe te videnticulata Lundell, Phytologia 61:66, 1986, nomen novum invalidum.
Type: COSTA RICA. ALAJUELA or PUNTARENAS: Cordillera de’ Tilaran, Flora of Monteverde,
comunidad, 1,400-1,500 m, 28 Nov 1976 (fl), Vv. Dryer 1010 (HOLOTYPE: F).

Ardisia parvipunctata (Lundell) See & Ricketson, comb. nov. Lcacorea

parvipunctata Lundell, Wrightia 7:274. 1984. Ardisia / ictata (Lundell) Lundell,
Phytologia 61:66. 1986, nomen valida ‘TYPE: MEXIC O. Oaxaca: Along high-
way 175, in the vicinity of La Galera, ca. 500 m S, 2.1 mi N of turn-off to Plama

Hid: ilgo, 9.6 mi S of Puente ies near village of Jalatengo, loose slope along
ravine above spring on highway, 1,340 m, 20 Jan ‘1979 (fr), 7) Croat 46150 (HOLO-
TYPE;

Ardisia samalana (Lundell) ae & Ricketson, comb. nov. /cacorea samalana
Lundell, Phytologia 57:45 5. Ardisia samalana (Lundell) Lundell, Phytologia
61:66. 1986, nomen es ca GUATEMALA. QUETZALTENANGO: Hot springs

along Rio Samala, below Zunil, 2,500 m, 24 Jan 1940 (fl), /. Steyermark 34962 (HOLOTYPE:
F).

jos

Ardisia steinii Pipoly X Ricketson, nom. nov. Auriculardisia ieee Lundell,
Wrightia 7:267. 1984, non Ardisia baruana Lundell, Wrightia 6:6 9. Ardisia
stemnit L. unde! Phytologia 61:67. 1986, nomen novum invalidum. a YPE: CON
CHIRIQUI: Volcan Baru, E slope on road from Boquete, 8 km W of Boquete (air dis-
tance), 8° 50' N, 82° 30' W, 1,900 m, 8 Jan 1983 (fr), B. Ste?n 1267 (HOLOTYPE: LL-
TEX; isoryepeE: MO).

Ardisia stevensii (Lundell) Pipoly & Ricketson, comb. nov. Gentlea stevensii
Lundell, Wrightia 6:97. 1979. Ardisia stevensti (Lundell) L ce Phytologia 61:67.
1986, nomen invalidum. Type: NICARAGUA. CHONTALES: ca. 2.8 km above (N
of) Cuapa, 12° 17' N, 85° | W, 400500 m, 21 Jan 1978 (fr), W. Stevens 6053

(HOLOTYPE: LL- TEX; tsorype: MQ).

Ardisia toroana (Lundell) Pipoly & Ricketson, comb. nov. Auriculardisia toroana
Lundell, Wrightia 7:273. 1984. Ardisia toroana (Lundell) Lundell, Phytologia 61:67.

1986, nomen invalidum. Type: PANAMA. Bocas DEL Toro: 15 km up the Changuinola

river to I.R.H.E. dam site No. 1, near campsite on trail to 1 idge NE of campsite,

1X;

800-900 ft [244-274 m], 12 Dec 1979 (fl), 7) Antonio 3079 (HOLOTYPE: LL-T
©).

=

ISOTYPE:

Ardisia triangula (Lundell) Pipoly & Ricketson, comb. nov. /eacorea triangula
Lundell, Phytologia 56:417. 1984. Ardisia triangula (Lundell) Lundell, Phytologia

PIroLy AND RICKETSON, New names and combinations in Myrsinaceae 515

61:67. 1984, nomen invalidum. Type: COSTA RICA. SAN José: Patarra, Cerro El
Espino, Alto Mata de Cafia-Azahar, 1,600—1,800 m, 13 Nov 1983 (fl, fr), G. Herrera
e I. Chacén 1579 (HoLoTYPE: LL-TEX; tsorypes: MO, NY).

Ardisia utleyi (Lundell) Pipoly & Ricketson, comb. nov. Icacorea utleyi Lundell,
Phytologia 57:452. 1985. Ardisia utleyi (Lundell) Lundell, Phytologia 61:68. 1986,
nomen invalidum. COSTA RICA. San José: 2-5 km SE of Higuito on Calle Tablazo
or 8-12 km SE of Desemparados, 1,800-1,900 m, 5 Sep 1975 (fl), /. Utley e K.
Utley 3023 (HOLOTYPE: F).

Ardisia warneri Pipoly & Ricketson, nom. nov. Leacorea brevipes Lundell, Phytologia
56:414. 1984, non Ardisia brevipes Lundell, Wrightia 3:97. 1964. Ardisia warneri
Lundell, Phytologia 61:68. 1986, nomen novum invalidum. Type: COSTA RICA
PUNTARENAS: Foothills of the Cordillera de Talamanca, around Tres Colinas, 1,800—
1,850 m, 20 Mar 1984 (fr), G. Davidse, G. Herrera Ch. & R. Warner 25680 (HOLO-
Type: LL-TEX; tsorypes: BM, MO, NY).

Ardisia wendtii (Lundell) Pipoly & Ricketson, comb. nov. /barraea wendtii
Lundell, Wrightia 7:46. 1982. Ardisia wendtii (Lundell) Lundell, Phytologia 61:68.
1986, nomen invalidum. Type: MEXICO. Veracruz: Municipio Minatitlan, 8.2
km al N de la terracerfa La Laguna-Rio Grande, sobre camino nuevo, a Ejido Belisario
Dominguez, el cual sale de la terraceria 14.7 km al E de La Laguna, 17° 22' N, 94
22' O, 130 m, 16 Jul 1980 (fl, fr), 77 Wendt, A. Villalobos, R. Lara, M. Navarrete &
I. Navarrete 2584 (HOLOTYPE: LL-TEX; tsotyres: F, MQ).

Finally, the following five taxa were originally described by Lundell (1984,
1987) as species of /cacorea, but our studies indicate they are members of
Ardisia subgenus Icacorea, to which we effect the transfers using the fol-
lowing new names and combinations.

ee ate (Lundell) ue & Ricketson, comb. nov. Jeacorea albipetala
dell, Phytologia 63:465. 1987. Tyre: PANAMA. Bocas DEL Toro: Region of
Ae Colorado, on trails from continental divide 7 mi. from Champi Camp, ca8"
35' N, 81° 45' W, 1,500 m, 12 Apr 1986 (fl), G. McPherson 8832 (HoLoTypeE: LL-
TEX; tsorypes: MEXU, MO, PMA).

Ardisia duripetala (Lundell) Pipoly & Ricketson, comb. n ‘orea duripetala
Lundell, Phytologia 63:464. 1987. ‘Type: ee caine S ie Be Ccn6
Horqueta, N of Boquete, 6,500 ft [1,981 m], 21 Jan 1971 (fl), R. Wilbur, J. Teeri er
R. Foster 13471 (HoLotyPe: LL-TEX).

Ardisia hatoana (Lundell) Pipoly & Ricketson, comb. nov. [cacorea hatoana
Lundell, Phytologia 63:464. 1987. Type: PANAMA. Cuiriqut: Near Las Lagunas,
W of El Hato del Volcan, ca. 1,400 m, 15 Jan 1970 (fl), R. Wilbur, R. Weaver, R
Foster e& M. Correa 11002 (HOLOTYPE: LL-TEX; tsotyPE: DUKE).

Ardisia lajana (Lundell) Pipoly & Ricketson, comb. nov. scacorea lajana Lundell,
Phytologia 63:465. 1987. Type: MEXICO. Veracruz: Municipio Zapata, La Laja,
entre Corral Falso y Pinaltepec, a 900 m de carretera Xalapa-Veracruz por camino
Corral Falso-Pinaltepec, 900 m, 8 Ene 1985 (fr), H. Herndndez y R. Torres 769 (HO-
LoryPe: LL-TEX; 1sorypes: CHAPA, ENCB, MEXU, MO)

Ardisia mesoamericana Pipoly & Ricketson, nom. nov. Jcacorea latisepala Lundell,
Phytologia 56:417. 1984, non Ardisia latisepala (Lundell) Pipoly & Ricketson, Sida
18: 513. 1998. (based on Auriculardisia latisepala Lundell, Wrightia 7: 269. 1984).

516 SIDA 18(2)
Type: PANAMA. Bocas bEL Toro: Cordillera de Talamanca, 8 airline km NW of

the peak of Cerro Echandi on the Costa Rican-Panamanian international border, 9°

5' 30" N, 82° 51' 30" W, 2,8002,900 m, 49 Mar 1984 (fl), G. Davidse, L. Gémez,
G. Herrera, C. Chacén e» A. Chacén 25279 (HOLOTYPE: LL-TEX; isorypes: BM, MO,
NY)

ACKNOWLEDGMENTS

We thank the Missouri Botanical Garden and the Flora Mesoamericana
Project, for funding that allowed J. Ricketson (MO) to travel to Fort Worth,
ce he was joined by J. Pipoly (BRIT) on visits to the C.L. Lundell Herbarium
(LL-TEX), housed at the University of Texas at Austin. Without access to
that critical collection, assembled by C.L. Lundell over a period of nearly
60 years, the present study would not have been possible. Flora Mesoamericana
and the Missouri Botanical Garden also provided funding for Pipoly to travel
to MO to work with Ricketson. We thank the curators of the herbaria cited
for loans of specimens. We are grateful to the staff of TEX, especially Billie
L. Turner, Tom Wendt, Carol Todzia, Beryl Simpson, and José Panero, for
their cooperation and hospitality. We are also grateful to those who have
been so instrumental in assisting us in our work, including K. Gandhi (GH),
Dan Nicolson (US), Gerrit and Jeany Davidse, Linda Oestry, Mary Bard,
and Catherine Mayo, (MO), Barney Lipscomb, Lindsay Woodruff, Marjorie
Weir and Jim Rivers (BRIT). Reviews of the manuscript by K. Gandhi and
G. Davidse, and meticulous copy editing by Barney Lipscomb, greatly improved
the presentation of the paper.

We also specifically thank Dave Boufford, K. Gandhi and Peter Stevens
(A, GH); R. Brummitt and K. Challis (K); M. Grayum and G. Davidse
(MQ); and D. Nicolson and W. Wagner (US) for reviewing the nomencla-
tural problems presented here and our proposed solution to them. While
we did not necessarily agree with all of their points of view, we thank them
for their interest, careful consideration of the subject, and the time they
spent explaining the points to us. Therefore, the opinions expressed in this
paper are exclusively our own. Our arguments were certainly made stron-
ger by the thought-provoking debates we shared with all the concerned parties.

REFERENCES
AUBLET, F, 1775. Histoire des plantes de la Guiane francoise. P-F. Didot. 2 vols.
Pica: E. 1940. A revision of the Hawaiian species of Myrsine (Suttonia, Rapanea) (Myrsinaceae).
Occ. Pap. Bishop Mus. 16:25—
LUNDELL, C.L. 1963. Studies of me Ammeiean Myrsinaceae- |. Wrightia 3:77-90.
. 4. Studies of the American Myrsinaceae- I. Wrightia 3: 97-114.
____. 1966. Myrsinaceae. In: P.C. Standley and L.O. at eds. Flora of Guatemala,
et Number 1. Fieldiana, Bot. 24 (8/2): 135—2(
. 1971. F ae 4 - Panama, Part VIII. Family 150. 7 nee Ann. Missouri Bot.
“Cat 58:285-.
sel, ae Myrsinaceae—lV. Phytologia 48:137-142.

PIPOLY AND RICKETSON, New names and combinations in Myrsinaceae Ble

. 1981b. Studies of American plants—XX. Phytologia 48:131—-136.
. 1981c. Neotropical Myrsinaceae—VI. Phytologia 49:341-354.
. 1982. Neotropical Myrsinaceae—VII. Wrightia 7:38-50.
. 1984, Neotropical Myrsinaceae—XV. Phytologia 56:413-419.
. 1985. Neotropical Myrsinaceae—XVI. Phytologia 57:449-452.
. 1986. Neotropical Myrsinaceae—XX. Phytologia 61:62-68.
. 1987. Neotropical Myrsinaceae—XXI. Phytologia 63:463-465.
Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pélanzenreich IV. 236 (Heft 9):1-437.
MORALES, J. F. 1997 [1 998]. Una nuevz a especie y las Myrsinaceae
de Costa Rica y Panama. Phytologia 83:109-112.
Pipoty, J. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia
(Myrsinaceae). Mem. New York Bot. Gard. 43:1-76.
. 1992. Notes on the genus Myrsine (Myrsinaceae) in Peru. Novon 2:392—407.
. 1996. Contributions toward a new flora of the Philippines: I. A synopsis of the
ae Myrsine (Myrsinaceae). Sida 17:115—162
cand J. Ricketson. 1998. A revision of the genus Ardisia subg. Graphardisia (Myrsinaceae).
Sida ns XXX—
RICKETSON, J. ai, PIROLY.. 199 7a nN latural notes and a synopsis of the genus Myrsine
Myrsinaceae) in poses Ie Sida 17:579-5.

ores and q svnonss of N f{
r

Stylogyne (Myrsinaceae).

~ Sida 17:591-598.
1997c. A. synopsis of the genus Gentlea (Myrsinaceae) and a key to the genera of

Nisin in Mesoamericana. Sida 17:697—707.

STEARN, W.T. 1969. A synopsis of Jamaican Myrsinaceae. Bull. Brit. Mus. Nat Hist. Bot.
Ser. 18:884-900

SWARTZ, O. 1788. Nova genera & species ori seu Prodromus. Holmiae [Stockholm],
Uppsalae [Uppsala], & Aboae [Abo]. 1-158.

Wacker, E.H. 1959. A revision of the ee of Taiwan. Quart. J. Taiwan Mus. 12:161-
194.

SIDA 18(2)
BOOK NOTICE

Wittiam A. Werper. 1997. King of Colorado Botany. Charles Christo-
pher Parry, 1823-1890. (ISBN 0 87081 431-1, hbk). University of
Colorado Press, PO. Box 849, Niwot, CO 80544. 303-530-5337.
$39.95. 183 Pp.

This book is about Charles Christopher Parry’s exploration of the southern Rocky Mountains

of Colorado. Weber introduces the life of Parry and his botanical collections that intro-

duced the Colorado flora to the world. “Over eighty new species of flowering plants were
named from Parry’s Colorado collections, and many more from his collections in Mexicc

ORs SSUES Two aes a te ya and Parryella, were dedicated to him.

an d the
ragalus parryi, Campanula parryi, hae

Some of the Colorado novelties, such a
parryi, Helianthella parryi, Oxytropis parryi, Pedicslets parry Primula parryi, and Tri
lium we are Colorado’s most well-known wildflowers.

The 183 page book on the “King of Colorado Botany” is broken up into the following:
Preface; pet ise pam an Introduction to The Collections and The Life of C.C. Parry;
1) Setting the State: The Parry Narratives; 2) Parry’s Colorado Expedition of 1861; 3)
The Parry, Hall, and Harbour Expedition of 1862; 4) Expedition of 1864; 5) Observa-
tions on Snow Line and Timberline; 6) Expedition of 1867; 7) Dedication of Gray’s and
Torrey’s Peaks, 1872; 8) Collections of 1873-1889; and 9) Cryptogamic Collections. These
chapters are followed by four appendixes: New Taxa Described from Parry’s Colorado
Collections, The Parr rbarium, Parry’s Dismissal as Botanist for the Department of
Agriculture, and Publications of C.C. Parry. References and Index follow the appendixes.

SIDA 18(2): 518. 1998

A NEW SPECIES OF FILMY FERN
(HYMENOPHYLLACEAE: PTERIDOPHYTA)
FROM SOUTH INDIA

C. ABDUL HAMEED

Department of Botany
M.ES.KVM. College
Valanchery
Kerala 676 552, INDIA

PV. MADHUSOODANAN

Department of Botany
University of Calicut
Kerala 673 635, INDIA

ABSTRACT

Crepidomanes indicum, a new species of filmy fern is described and illustrated from southern

Western Ghats of India.
RESUMEN

Se describe e ilustra Crepidomanes indicum, una nueva especie de helecho membranoso
de los Ghats del suroeste de la India

Crepidomanes is an Old World genus of more than a dozen species rang-
ing from East African islands to Japan and Polynesia (Tagawa & Iwatsuki
1979). The name derived from two Greek words viz., Krepis = slipper +
manes = cup, alluding to the shape of the indusium (Stewart et al. 1983).
The presence of striae and a submarginal pseudovein are the conspicuous
diagnostic characters of this genus. Beddome (1883) has reported three species
now known as C. kurzii (Bedd.) Tagawa & K. Iwats., C. intramarginale
(Hook. & Grev.) Copel. and C. bipunctatum (Poir.) Copel. from South India.
Later, Nair et al. (1992) and Manickam & Irudayaraj (1992) reported the
occurrence of C. latealatum (Bosch) Copel. and C. plicatum (Bosch) Ching.
respectively from the South India. Recently Madhusoodanan and Hameed
(1997, 1998) reported C. christii (Copel.) Copel. and C. agasthianum
Madhusoodanan & Hameed from the dense evergreen forests of western
Ghats, South India.

While revising the genus Crepidomanes of India, the authors collected a
very small filmy fern from the mature, dense, tropical evergreen forest of
Thommankuthu in the foothills of the Western Ghats mountains in the

SIDA 18(2): 519-522. 1998

520 SIDA 18(2)

Idukki district of Kerala state, S. India (alt. 700 m; annual rainfall 3000
mm; temp 20-35°C). The plant shows a high degree of differentiation from
the species previously described under the genus Crepidomanes (Beddome
1863, 1866, 1883; Copeland 1938; Devol 1975; Holttum 1954; Iwatsuki
1984, 1985; Jinn-Lai & Wang-Chueng 1994; Sledge 1968). It differs from
other species in its smaller size, dark green, evenly pinnatifid fronds having
the rachis-wing and segements with an entire margin; thin submarginal veins
and cupular indusia with highly dilated mouths. The plant is described here
as Crepidomanes indicum.

se ea indicum Hameed & Madhusoodanan, sp. nov. (Fig. 1) ‘Tyr

NDIA. Kerala: Idukki Dt.: Thommankuthu water fall, 600 m, 1 Sep 1996, C. A
See CU 34880 (HOLOTYPE: CALI; isoTypes: K, MH).

Crepidomani agasthiano ffinis, sed frondibus atro-viridibus profunde, regularite er pinnatifidis
segmentis aequalibus, margine recto, apice vad inato, venula fala intra- — ali
tenui, cellulis marginalibus latis unistratis contents srepletis differt. S 2-8 terminales,
involucre cupuliformes, latiore quam longiore, parti ke ae

ore maxime dilatato.

Rhizome filiform, long, creeping, profusely branched, less than 0.25 mm

thick, bearing thick walled, dark brown, unicellular rhizoids, 0.5—-1.5 mm
long. Fronds scattered on rhizome at intervals of 2—4 mm; stipes 3—5 mm
long, covered with profuse hairs like those of rhizomes (but smaller in size).
Laminae regularly pinnatifid (pinnate-pinnatifid or rarely bipinnatifid) 2—
7 mm long, thin, membranous. Pinnae 2 or 3-lobed, segments ca. 0.6—0.7
mm wide, margins entire, single-celled in thickness with shallowly emar-
ginate apex. Laminal segments each with a single vein ending at the apex,
arising alternately from the midrib, clothed with sparse, clavate trichomes;
intramarginal pseudoveins thin and continuous, separated from the mar-
gin by a single row of comparatively broad, marginal cells; other false veins
absent. Sori 2-8, embedded at the apices of distal segments, involucres cup-
shaped, broader than long, ca. 1 x 1.2 mm, involucral halves over-full, mouths
highly, dilated but without lips. Receptacle filiform, included. Sporangia
subsessile, spherical with an oblique annulus composed of 16-21 indurated
cells without any thin walled cells; spores tetrahedral, trilete, greenish, perineless,
30—43 mm in diameter, with a sparsely tuberculate exine.

Distribution.— Crepidomanes indicum is not very common and so far
known only from Athirapally forest of Trichur district (ca. 200 km south),
in addition to the type locality.

Ecology.—Crepidomanes indicum was found growing epiphytically on the
trunks of the large trees (about 15-20 m height) in the dense evergreen
riparian forests. The black wiry rhizome forms a prominent network on
the bark of trees about 1.5 m above the ground. The species is confined to
the trees growing very near to the streams.

Notes.—Crepidomanes indicum resembles C. agasthianum Madhusoodanan

HAMEED AND MaDHUusooDANAaN, A new species of Crepidomanes 521

ae,
S\ \Z-
y.

;
/ os

FiG. 1. Crepidomanes indicum. A. Habit; B. Sporangium; C. Spores; D. Hairs on rhizome;
E. Tip of fertile frond with sori with sporangia; left sorus showing view with one half of
the indusium removed; F. Trichomes on veins; G. Details of frond margin with intramarginal
pseudovein.

& Hameed in appearence. It shows an affinity to Microtrichomanes nitidulum
(Bosch) Copel. in indusium shape and to Microgonium henzaianum (Par-
ish ex Hook.) Copel. in the shape of its involucral halves. It differs from C.
agasthianum in having pinnatifid, dark green fronds with entire margined
laminae and cup-shaped, broader than long indusia with dilated but not
spreading mouths. Crepidomanes agasthianum has pale green, digitate, nar-
rowly segmented fronds with undulate margins and cupular sori with nar-

—_—

522 Stipa 18(2)

row mouths subtended by branches of costa running into two long over-
topping segments. Sori are characteristic in their dilation of the involucral
mouths as well as in the overfullness of the involucral halves, which brings
about the entire length of receptacle with sporangia visible from the upper
view. These characters, along with the extreme dwarfness of the plant, dif-
ferentiate this new taxon from C. kurzit, which is luxuriantly growing in
the type locality.

Pararypes: INDIA. Kerala. Idukki Dt.: Thommankuthu stream, near Thodupuzha, 750
m, 27 Dec 1996, C.A. Hameed CU 34810 (CALI); Thommankuthu forest, 750 m, 13
Dec 1997, C.A. Hameed CU 34853 (CALI). Trichur Dt.: Athirapally forest, 1 km above
Athirapally waterfall, 800m, 21 Dec 1997, CA. Hameed, CU 34859 (CALI); Athirapally
waterfall, 21 Dec 1997, C.A. Hameed CU 34864 (CALI).

ACKNOWLEDGMENTS
The authors are grateful to J.F. Veldkamp, Rijksherbarium, Leiden, The
Netherlands, for the Latin diagnosis.
REFERENCES
Beppome, R.H. 1863. The ferns of southern India and Ceylon. Gantz. Bros., Madras.
Beppomg, R.H. 1866. The ferns of British India. Gantz Bros., Ma
Beppome, R.H. 1883. A handbook to the ferns of British India, Ceylon and Malay Pen-
insula. Thaiker Spink & Co., Calcutta.
Devot, C.E. 1975. Hymenophyllaceae. . H. Li, T. Liu, T. Huang, T. Koyama and CLE.
Devol, eds. Flora of Taiwan. Pp. 97-12
Hourrum, R.E. 1954. A revised flora of M: ze aya-Il. Ferns of Malaya. Govt. Press, Singapore.
IWATSUKI, K. 1984, Studies in the systematics of filmy ferns VII. A aes of classification
ased chiefly on the Asiatic species. Acta Phytotax. Geobot. 3:165—179.
fara KI, K. 1985. The Hymenophyllaceae of Asia, excluding Malesia. a Fac. Sci. Univ.
ae. Sect. 3, Bot. 13:501-551.
Junn-Lat, T. and S. WANG-CHEUNG. 1994. Be ras In Huang, Tseng-Ching,
ed. Flora of Taiwan, Taipei. Second Edition. 1:99—
MaADHUSOODANAN, P.V. and C.A. HAMEED. 1997. mes christit (Copel.) Copel.
enim new filmy fern species from India. Indian J. Forest. 20:190—
OD.

ee JSOODANAN, P.V. and C.A. HAMEED. 1998. Crepidomanes agasthianum, a new filmy
fern species (Hymenophyllaceae - Pteridophyta) from India. Nordic. J. Bot. 18:169—
170.

ManickaM, V.S. and V. IRupayaraj. 1992. Pteridophyte flora of the Western ghats—South
India. New Delhi.

Nair, N.C., S.R. GHosn, and P. BHARGAVAN. 1992. Fern-allies and ferns of Kerala—Part
IV. J. Econ. noe Bot. 16:449-53.

SLEDGE, W.A. 1968. Hymenophyllaceae of Ceylon. J. Linn. Soc., Bot. 60:289-308.

STEWART, R. D.M. JOHNSON and J.T. MickeL. 1983. Pteridophyte genera. The mean-
ing of their names. Fiddlehead Forum. 10:21—36.

Tacawa, M. and K. Iwarsuki. 1979. Flora of Thailand. Thailand Institute of Scientific
and Technological Research. Bangkok, Pp. 68-99.

A SECOND SPECIES OF ORITROPHIUM
(ASTERACEAE: ASTEREAE) FROM MEXICO

GUY L. NESOM

BONAP-NCBG
Coker Hall CB 3280
University of North Carolina,
Chapel Hill, NC 27599-3280, U.S.A.

ABSTRACT

Oritrophium durangense Nesom, sp. nov., is described from the Sierra Madre of south-
ern Durango, Mexico. It is the second species of the genus known from Mexico, following
another recently described from the northeastern slopes of Pico de Orizaba in Veracruz,
about 1000 kilometers northwest of the Durango locality. The remainder of the species of
Oritrophium occur in montane habitats of northern South America.

RESUMEN

Se describe CTD Cee Nesom, sp. nov., de la Sierra eee del sur de Durney,
México. E Mexico
descrita de las ders norcre del ies de Orizaba en Vereen unos 1000 kilémetros al
noroeste de la localidad de Durango. Las otras especies de Oritrophium se dan en habitats
del norte de América del Sur.

Identification and review of specimens of Asteraceae tribe Astereae in
LL, TEX have brought to light a collection of the genus Oritrophium (Kunth)
Cuatrec. from near the crest of the Sierra Madre in southern Durango, Mexico.
These plants do not correspond to the single previously described species
of Oritrophium in Mexico (Nesom 1992) or any other species of the genus
from its center of diversity in South America (Aristeguieta 1964; Cuatrecasas
1961, 1969, 1997). The plants from Durango are described here as a new
species.

Oritrophium durangense Nesom, sp. nov. (Fig. 1). Tyre: MEXICO. Duranco:
Mpio. El Salto, 4 km de la desv. hacia San Miguel de Cruces, bosque de pino-encino
principalmente, suelo profundo, con piedra suelto, 2200 m; herbacea de 15-25 cm
de alto, flores amarillas, en cabezuela, abundante; 6 Jul 1982, R. Hernandez 7676
con P Tenorio (HOLOTYPE: TEX!). The specimen was identified (on the distributed
label) as “Pionocarpus [= lostephane] af. madrensis (Wats.) Blake.”

A eae dad erable Nesom differt caudice tenui foliis basalibus paucis, foliis integris,
foliis t is longioribus, et floribus radii paucioribus limbis multo brevioribus.

Perennial herbs from a short, thick (ca. 8 mm) fibrous-rooted rhizome,
producing an erect, unbranched stem and a cluster of erect-ascending basal
Sipa 18(2): 523-526. 1998

Sipa 18(2

OE,

Fic. 1. Habit illustration of Oritrophium durangense, from the type collection.

Nesom, A second species of Oritrophium 525

leaves. Stems 12—16(—25) cm tall, purplish, minutely pubescent with a mixture
of Type B and Type C trichomes (see Nesom 1976 for illustrations), most
densely so near the heads. Leaves basal (largest) and cauline (reduced in
size); basal leaves linear, 1-nerved, entire, flat, somewhat succulent, 7-14
cm long, 3-6 mm wide, gradually tapered to a long, sharp point, glabrous
except near the base, where long, thin, cobwebby hairs (Type B) become
copious and conspicuous near the leaf insertion; cauline leaves reduced in
size from the basal, linear-lanceolate, 10-14 mm long, 2—2.5 mm wide at
the base, subclasping but not at all auriculate, 7-11 per stem, relatively even-
sized up the stem, with internodes 1-3 cm long, most closely spaced to-
ward the base. Heads radiate, turbinate, the involucres 10—14 mm wide;
phyllaries thin-herbaceous, without conspicuous venation, purple on the
margins and apex, nearly flat, oblong-lanceolate, mostly with acute apices,
slightly lacerate-fringed on the distal margins, in 3(—4) graduated series,
the innermost 8-10 mm long, ca. 2 mm wide, the outermost 1/2—2/3 as
long as the inner; receptacles flat, epaleate. Ray flowers pistillate, ca. 10—
15 in a single series, the corollas 6—7 mm long, the limbs 3—5 mm long,
barely exserted from the head, white or creamy, 1.8 mm wide, apically coiling,
densely invested with viscid, unicellular hairs around the tube-limb junc-
tion, the tube ca 2 mm long; disc flowers functionally staminate, with regular
corollas, 5 mm long, yellow, narrowly tubular-funnelform, the lobes trian-
gular, 1 mm long, the style branches linear-lanceolate, 1.8 mm long, densely
long-papillate, without stigmatic lines. Ray achenes fertile, densely sericeous,
eglandular, 5-nerved, ca. 5 mm long (not completely mature), narrowly
oblong, with pappus of numerous barbellate bristles 4-5 mm long in 1-2
series; disc ovaries sterile, with pappus bristles nearly smooth, equalling the
disc corolla length.

The new species is known only from the type collection. The type sheet
apparently bears two plants (one without the root/caudex system) and four
basal leaves disconnected from a plant. The illustration “attaches” several
of these leaves to the more complete plant. It is likely that a single plant
may produce several flowering stems, as is characteristic of other species of
the genus.

The plants of Oritrophium durangense are simple in habit but distinct in
appearance, producing a few, linear, basal leaves, a single, monocephalous
stem with a few, small, nearly bracteate cauline leaves, and a relatively small
head. The ray corollas are white and apparently barely exserted from the
involucre, perhaps accounting for why the collectors described the flowers
as yellow (the color of the disc corollas). Still, it seems unusual that the
species is currently known only from a single collection, since numerous
botanists have passed through the area and made collections. The collec-
tors of O. durangense noted that it was “abundante.”

526 Sipa 18(2)

These plants are unequivocally identified as the genus Oritrophium by a
distinctive set of features: monocephalous herbs producing a basal rosette
of elongate leaves, white rays, functionally staminate disc flowers (with sterile
ovaries and associated features of the style branches), phyllaries with in-
conspicuous venation, copiously elaborated, thin, nearly floccose white hairs
(Type B trichomes) in the axils of the basal leaves, and a lack of long, uniseriate,
thick-walled trichomes (Type A trichomes). The new species is perhaps related
to O. orizabense and a small group of Venezuelan species (Nesom 1992),
but this is speculative. The two Mexican species are different enough from
each other that even a hypothesis of sister relationship between them is obscure.

The new species differs from Oritrophium orizabense in its relatively thin
caudex/rhizome (8 mm vs. 7—15 mm) with few basal leaves (vs. dense clus-
ter of basal leaves), entire (vs. serrate) leaves, longer leaves (basal 7-14 cm
vs 1.5—4.0 cm, cauline 10-14 mm vs. 4-6 mm), and fewer ray flowers (10—
15 vs. 21-37) with much shorter limbs (3—5 mm vs. 7-10 mm). While the
habitat of O. orizabense is a cliff face, that of O. durangense is in soil (“suelo
profundo”), probably in a relatively flat area, with loose rocks (“con piedra
suelto”). The habitat of O. orizabense apparently is more like that of most
of the South American species, which generally grow “in wet places, the
marshy and swampy ground of the paramos, locations that are visited by
birds looking for water and seeds” (Cuatrecasas 1997, p. 289).

Oritrophium orizabense is disjunct from the main part of the genus (from
the closest point of distribution in Venezuela) by about 1500 kilometers.
The separation between O. durangense and O. orizabense is about 1000 kilometers,
with the latter the midpoint in more or less a straight line, making the dis-
tance between the Durango locality and those in South America about 2500
kilometers.

ACKNOWLEDGMENTS
I am grateful to the staff of LL, TEX for accomodating a recent visit there,

Harold Robinson for providing a copy of the recent synopsis by Cuatrecasas,
and John Pruski and Tim Lowrey for comments on the manuscript.

REFERENCES
ARISTEGUIETA, L. 1964. Compositae. Flora de Venezuela 10:1—483.
Cuatrecasas, J. 1961. Notas sobre Astereas Andinas. Ciencia [Mexico] 21:21-32.
Cuatrecasas, J. 1969. Prima flora Colombiana. 3. Compositae—Astereae. Webbia 24:1-

335.
CUuATRECASAS, J. 1997. Synopsis of the neotropical genus Oritrophium (Asteraceae: Astereae)
BioLlania, Edicion Especial 6:287-303.
Nesom, G.L. 1976. A new species of Evigeron (Asteraceae) and its relatives in southwest-
Utah. Brittonia 28:263—272.
Nesom, G.L. 1992. Oritrophium orizabense (Asteraceae: Astereae), a new species a and the
first report of the genus from North America. Phytologia 73:338-344.

UNIQUENESS OF THE ENDANGERED FLORIDA
SEMAPHORE CACTUS (OPUNTIA CORALLICOLA)
DANIEL FE. AUSTIN and DAVID M. BINNINGER

Department of Biological Sciences
Florida Atlantic University
Boca Raton, FL 33431, U.S.A.

DONALD J. PINKAVA
Department of Botany

Arizona State University

Tempe, AZ 35878, U.S.A.

ABSTRACT

Morphological analysis led to the conclusion that Florida Semaphore Cactus is a dis-
tinct species. That conclusion is congruent with genetic studies reported elsewhere. Our
plants are properly called Opuntia corallicola (Small) Werdermann (O. subgenus Consolea).
The Florida plants are not O. spinosissima, as recorded in recent literature. Indee
Florida plants are probably more closely related to O. hag and O. millspaughii han
to O. spinosissima. Details of the Florida species, and a key to similar and confusing Car-
ibbean relatives are provided. Because of the limited number of individuals remaining in
the wild, the threat from the alien pest moth Cactoblastis cactorum, the low genetic diver-
sity within Florida, and recent damage by Hurricane Georges, the Semaphore Cactus may
be the most endangered plant in the United States

RESUMEN

El estudio morfoldgico nos lleva a la conclusién de que el cactus semaforo de Florida es
una especie distinta. Esta conclusién esta de acuerdo con los estudios genéticos publicados
en otros lugares. Las plantas son propiamente llamadas Opuntia corallicola (Small) Werdermann
(O. subgénero Consolea). Las plantas de Florida no son O. spinosissima, tal como fue publicado
en la literatura reciente. Es mas, las plantas de Florida probablemente son mas cercanas a
O. rubescens y O. millspaughii que a O. spinosissima. Se ofrecen detalles de la especie de
Florida y una clave taxonédmica de las especies emparentadas y confusas del area del Car-
ibe. Dado el numero limitado de individuos que quedan en estado salvaje, la amenaza de
la polilla aléctona Cactoblastis cactorum, la poca ee genética ante de Florid: as a el
reciente impacto del huracan Georges, q la plar

de los Estados Unidos de Norte ee (EEUU).

In 1930, John K. Small described a cactus from the Florida Keys as a
new species that he called Consolea corallicola. Very little was known about
the species in the 1930s and in the subsequent decades until it became a
candidate for endangerment in the 1970s under the name O. spinosissima.

Sipa 18(2): 527-534. 1998

528 SIDA 18(2)

Cactus hobbyists were thought to have eliminated the species from the state
in the late 1970s. Even George Avery, a student of the Keys flora for 25
years at the time, could not locate the oe because they no longer grew
where he had known them (Austin et al. 1980; Avery 1981). Somewhat
later Ann Williams (in litt. 1986) found plants in the lower Keys and brought
them to everyone’s attention. Eventually the land on which the Semaphore
Cactus grew was purchased by The Nature Conservancy and it is now managed
as a preserve (Gordon & Kubisiak 1998; Negrén-Ortiz 1998).

Although the Florida Semaphore Cactus rarely appears in the literature,
there has been a raging controversy behind the scenes about what species it
really is. Long and Lakela (1971) seem to be responsible for first calling the
plants Opuntia spinosissima Miller, and they recorded it as endemic to Florida
even though that name was given originally to Jamaican plants (Britton &
Rose 1937). Opuntia spinosissima was picked up by those compiling the
“Smithsonian List” of endangered species from the 1970s and that name
continued to be used in subsequent publications (e.g., Benson 1982; Palmer
1984; Wunderlin 1998; Negrén-Ortiz 1998).

In the early 1980s, unpublished letters were being exchanged by Rich-
ard Howard of Harvard University and George Avery of Miami. Swayed
by Howard and Touw’s (1982) comments on the Lesser Antilles plants, Avery
decided that the Florida Keys plants must be Opuntia rubescens Salm-Dyck
ex DC. (G. N. Avery, in litt.).

This was the state of affairs in 1990 when Doria Gordon (The Nature
Conservancy, Gainesville), who was concerned about their plants on the
preserve, brought the problem to our attention. We present here the first
results of our studies. Other studies examine the genetic relationships of
these endangered plants and their relatives (Dougherty 1996; Gordon &
Kubisiak 1998).

‘To discover the evolutionary and nomenclatural status of the Florida plants,
it was first necessary to determine what taxa have been considered related
to those plants. Various authors have considered this alliance at different
ranks. Some consider Consolea Lemaire a distinct genus (Areces 1996). We
consider it a subgenus, e.g., Opuntia subg. Consolea (Lemaire) A. Burger.

Methods. Survey of the literature revealed nine nominate species. Areces
(1996) has excluded O. bahamana from them and we concur (Fig. 1). We
were able to obtain live specimens of six of the taxa. Live material of these
six remains in the collections at Fairchild Tropical Garden. Duplicates have
been sent to the University of Arizona and Desert Botanical Garden, Tempe.
From the living material available, and the literature records of the other
species, we created a data matrix of morphological traits that may be used
for recognizing these various plants. We present here only those six species
most likely to be confused with the Florida plants (Table 1).

AusTIN, et al., Uniqueness of Opuntia corallicola 529
O. corallicola ~- dy’ = o O. millspaughii
O. macracantha OY aire sash

O. moniliformis
| O. millspaughii

Pies
| S eee
c
QO. millspaughit _
~~
4 ~ a es
- QO. spinosissima O. falcata O. rubescens
N O. moniliformis v

O. rubescens X

VEMEZUELA

Fic. 1. Distribution of the species in Opuntia subgenus Consolea. This subgenus is con-
fined to the Caribbean.

Results and Conclusions. Confusion of O. corallicola with O. spinosissima
apparently resulted from previous studies not having examined living plants
(Austin and Pinkava 1991). Although names are not fixed by typification
in O. spinosissima and O. rubescens, we are applying them consistently with
historical usage. Moreover, the morphotypes have distinct ranges (Table 1)
consistent with historical name application, and with the usage by Areces
(1996).

As stated by Britton and Rose (1937), O. spinosissima is endemic to Ja-
maica. Although O. rubescens is widespread in the upper Antilles (Fig. 1),
it too is distinct from the Florida plants. In addition to morphological traits,
O. rubescens has 2n=132 chromosomes in the living material we examined.
Previous reports have given Opuntia rubescens as 2n=22 and 132 (Spencer
1955; Katagiri 1952; Yuasa et al. 1973). We do not know if these reports
represent different chromosome races or misidentifications. At least the numbers
are consistent with polyploid series known within the genus (Pinkava et al.
1985).

Analysis of morphology indicates that the Florida population of Sema-
phore Cactus, O. corallicola, is related to O. millspaughii, O. rubescens, and

TABLE 1. Comparative morphology of Caribbean species confused with O. cora/licola.

Traits corallicola Spimosissima ribescens millspaughii nashit moniliformts
Distribution Florida Keys Jamaica Hispaniola & = Cuba and the Bahamas Cuba

Puerto Rico Bahamas to

to Guadelupe Caymans
Habitat mesic Xeric xeric Xeric X€TIC Xeric
Height 3.5m 1-3.5 m 5 +4m 1-3.5 m =5m
Are not reticulate reticulate not reticulate reticulate not reticulate reticulate
Areole distance 5 cm 1-1.5 cm 1-1.5 cm 1-1.5 cm 3 1-1.5 cm
Areole level su raised raised raised & pitted raised & pitted raised
Pad shape elliptic oblong-linear oblong-linear lanceolate to oblong-linear oblong-linear

Joint length

Spine color

Spine location
Maximum spine number
Spine shape

Spine length

Spine direction

soo

vericarpel length
Pruit

Chromosomes

gray or white
entire surface
1-5

acicular
7-12 cm
deflexed
acute

10-15 mm

proliferous

to oblong
3-5 dm
straw

entire surface
1-5

acicular
7-12 cm
deflexed
rounded
10-15 mm

non-proliferous

to oblong

entire surface
subulate
7-12 cm
spreading
acute

10-15 mm

proliferous
132

es: Za

lanceolate-oblong
3-5 dm

gray or white
marginal

acicular

>12cm

deflexed

acut

=5 mm

3—4 cm

non-proliferous

to oblong
=10 dm
gray or white

entire surface

acute
10-15 mm

3—4 cm
proliferous
2n=?

to oblong
1-3 dm

gray or white
marginal

1-5

subulate
7-12 cm
spreading
rounded
10-15 mm

5-6 cm
non-proliferous

OS

(Z)81 Vals

AUSTIN, et al., Uniqueness of Opuntia corallicola 531

the Jamaican endemic O. spinosissima (Austin and Pinkava 1991). Chloro-
plast DNA study (Dougherty 1996) supports that conclusion. The mor-
phological data (Table 1) indicates more similarities between O. millspaughii
and O. corallicola than between the others. RAPD data (Gordon & Kubisiak
1998) also indicate that O. corallicola is more closely related to O. millspaughii
(Cayman Brac plants of Gordon & Kubisiak 1998) than to O. spinosissima.

As the Cayman Brac plants are a species (O. millspaughii) that ranges
geographically closer to Florida than Jamaica, the morphological and ge-
netic results are consistent with biogeographic predictions (Brown 1998).
More exact relationships among the various species within Opuntia subg.
Consolea, and between those species and the remainder of the genus will be
reported by Areces (New York Botanical Garden).

We conclude that Small (1930) was correct when he proposed the Florida
Semaphore cacti as distinct. While this uniqueness was confused by use of
the name O. spinosissima in recent literature, available data indicate that
the Florida and Jamaica plants are markedly different species. The Jamaica
plants are morphologically unique (Table 1) and endemic to that island, as
originally recorded by Britton and Rose (1937).

TAXONOMY

The complete taxonomy of the group Consolea is being revised by Areces
(1996). Here, only the Florida taxon will be addressed.

Opuntia corallicola (Small) Bakeberg, Neue Kakteen, Jagden, Arten, Kultur.
Gartenbau- Verlag Trowitzsch and Shon, Frankfurt (Oder)-Berlin. 1931.
Consolea corallicola Small, Addisonia 15:25—26, pl. 493. 1930. Type: FLORIDA.
Monroe Co.: Big Pine Key, 1919, Small (syntype: NY!); Key Largo, Small (syNTYPE:
NY!). No lectotype is chosen in deference to Areces (in prep.).

Opuntia spinosissima sensu authors, non Miller (1768).

Description: Shrub or small tree 1—3.5 m tall. Trunk nearly cylindrical,
0.5—2.5 m long, reaching 3—4 cm in diameter. Larger terminal joints light
green, standing mostly ascending, all flattened, ca. 1 cm thick, mostly el-
liptic, but ranging to elongate or asymmetrical, 1-3 dm long. Areoles el-
liptic, typically 1-1.5 cm apart. Spines numerous, in all areoles or some
joints nearly spineless, gray or white, turning brown with age, 2—3(—4) per
areole, spreading and deflexed, the longer 7—12 cm, basally 0.25—0.5 mm
in diameter, acicular, nearly circular in cross section, twisted, somewhat barbed.
Glochids yellow, abundant, 1-1.5 mm long. Leaves rudimentary, small,
deciduous, scale-like. Flowers 1.2—2.5 cm in diameter. Sepals green, ovate-
deltoid, 3-6 mm long, mostly acute. Petals orange-yellow, turning red shortly
after opening, broadly ovate-acute, mucronate, entire. Filaments yellow, 6
mm long; anthers yellow, 0.5 mm long. Styles 6—7.5 mm long, ca 0.5 mm
in diameter; stigmas 5, thick, mostly 1.5 mm long. Ovary in anthesis spiny,

532 Sipa 18(2)

flattened. Fruits when produced turning yellow, 5—6 cm long, proliferous;
seeds few, circular and flattened, irregular, the edges cristate, the sides puberulent,
6-8 mm in diameter.

Range.—Known from 12 plants with vegetative seedlings on The Na-
ture Conservancy preserve in the Florida Keys; also a few individuals that
have been brought into cultivation from sites in the keys, both known and
unknown. Formerly known from Key Largo and Big Pine Key.

IHustrations.—A color plate 493 was published by Small (1930).

Comments.—Britton and Rose (1937) provided a black and white plate
of O. spinosissima with a longitudinal section of the flower and ovary showing
the nectar chamber typical of subgenus Consolea. That chamber is also found
in Nopalea. Areces (1996:230) has given superlative diagnostic sketches of
the flowers of four other species in subgenus Consolea: O. macracantha, O.
moniliformis, O. nashit, and O, millspaughii. The flowers of O. corallicola
are most similar to those of O. mi//spanghii, but these two differ in several
other traits (Table 1)

The nectary chamber, the unjointed central woody axis, and distinctive
pollen (Leuenberger 1976), are among traits that make Conso/ea distinc-
tive, regardless of the rank it is given (Areces 1996).

KEY TO SOME CONFUSING CARIBBEAN OPUNTIA
1. Spines straw-colorec
2. Plants 1—3.5 m tall. Areoles reticulate. Joints 3—5 dm long. Spines 1-
5, acicular, deflexed. Petal apex rounded. Pericarpel 7-8 cm long. Fruits
O. spinosissima

not proliferous
2. Plants 5-10 m tall. Areoles not reticulate. Joints 1-3 dm long. Spines

5-9, subulate, spreading. Petal apex acute. Pericarpel 5—6 cm long.

Fruits ean O. rubescens

. Spines gray or white (brownish with age).
: | y
piceeuate Spines >12 cm long, mostly marginal. Fruits non-
prolife rous. O. millspaughii

3, Areoles not reticulate. Spines 3-12 cm long, over entire surface. Fruits
proliferous.
4. Areoles sunken, 1-1.5 cm apart. Joints 1-3 dm long. Pericarpels

5-6 cm lot O. corallicola
4, ee ie oo and pitted, 1.5—-3 cm apart. Joints = 10 dm long.
Pericarpels 3—4 cm long. O. nashii

The future. Current findings indicate that the Florida plants are an ex-
ceptionally rare and endangered endemic species. Presently O. corallicola is
known from 12 plants on one small land parcel that is owned and pro-
tected by The Nature Conservancy. That organization has formulated and
implemented an informal recovery plan for this endangered species, but
that procedure is complicated by recent events.

The presence of the exotic moth Cactoblastis cactorum has initiated a series
of problems with those plants (Pemberton 1995). This moth has substan-

AusTIN, et al., Uniqueness of Opuntia corallicola 533

tially reduced the abundance of Opuntia in the Keys and attacked the Semaphore
cacti in the past. Those attacks resulted in most of the remaining plants
being placed within screened “cages” in 1990. Although the cacti grow in
partial shade conditions, the exclosures may have altered the light quality
reaching the plants and caused growth anomalies such as etiolation. No
data are available on these characters. Moreover, the cages exclude any po-
tential pollinators of the cacti, precluding potential allogamous sexual re-
production. Future survival with this insect pest is problematical for the
Florida plants in spite of the dubious declining infestation trends predicted
by Johnson and Stiling (1998). That short-term study, terminated in 1993,
did not include the Florida Atlantic University (FAU) campus (Pierce 1995),
nor did it address the more recent (1996-1998) upswing in infestation in
southern Florida. Cultivated plants on the FAU campus were destroyed during
1997 by Cactoblastis, after the native population of O. stricta was decimated.

A second more recent event has led to another problem for the plants.
Hurricane Georges swept through the Florida Keys in September of 1998.
The exclosures were removed when the Hurricane was approaching and
have not been replaced. With that event, storm surge swept over the low
limestone keys and deposited ocean water. That salty deposit was not im-
mediately diluted by subsequent rainfall, and its long-term impact on the
plants is unknown. Also, the main stems on two of the individuals were
broken. On others the stems remained erect, but the pads were removed
(D. Gordon, pers. comm., 13 Oct 1998). Smaller propagules at the bases
of the cacti remain. The full impact of the hurricane on these plants is still
being evaluated.

ACKNOWLEDGMENTS

This research was supported by U.S. Fish and Wildlife Service Contracts
No. 14-1608004-79-106 and FAU 02-1140-090 (D. E Austin investiga-
tor), 41910-2-0174 and 41910-3-0359 (D. FE Austin and D. M. Binninger
co-investigators). We thank Sandra K. Austin; Doria R. Gordon and Randy
‘Tate (The Nature Conservancy); Les Landrum and Jon Rebman (Arizona
State University); R. PR Wunderlin (University of South Florida); Vivian
Negron-Ortiz (Miami University of Ohio); Alberto E. Areces-Mallea (New
York Botanical Garden), and anonymous reviewers for comments on drafts
of the mss. Santiago Pérez (FAU, Biology) kindly provided the Spanish

translation.

REFERENCES

Areces-M., A. 1996. New taxa of Consolea Lemaire (Cactaceae: Opuntioideae) from Cuba.
Brittonia 48:224—236.
AusTIN, D.F, P. Krauss, J.L. Jones, C.E. Nauman, and B.E. Tatje. 1980. Endangered species

534 Sipa 18(2)

of southern Florida. Report to U.S. Fish and Wildlife Service, Office of Endangered
ae Atlanta, GA. Contract No. 14-1600004-79-106.

—-and D.]J. Pinkava. 1991. Preliminary summary of the taxonomic status of the Florida
Semaphore Cactus. report to U.S. Fish and Wildlife Service, Office of Endangered Spe-
cies, Jacksonville, FL.

Avery, G. 1981. A history of the Cactaceae in the southeastern United States. Cac. Succ.
J. (Los Angeles) 53:177-182.

BAKEBERG, C. 1931. Neue Kakteen, Jagden, Arten, Kultur. Gartenbau-Verlag Trowitzsch
and Shon, Frankfurt (Oder)-Berlin.

BENSON, L. 1982. The cacti of the United States and Canada. Stanford University Press,
Stanford, CA.

Brirron, N.L. and J.N. Rose. 1937. The Cactaceae. Descriptions and illustrations of plants

of the Cactus Family. Dover Publ. Inc., New York.

Brown, J. H. AND M. V. LoMOLINO. 1998. Biogeography. Second ed. Sinauer Associates,
Inc., Sunderland, MA

DoucGuerry, C. J. 1996. Nel DNA sequences of the endangered Florida Semaphore Cactus.
M.S. Thesis, Florida Atlantic University.

Gorbon, D.R. and T.L. Kusisiak. 1998. RAPD oe of the last populations of a likely

Florida Keys endemic cactus. Florida Sci. 61:203-21

Howarp, R.A. and M. Touw. 1982, Opuntia species in ae Lesser Antilles. Cac. Suce. J.
(Los Angeles) 54:170- 179.

KaTiciri, $. 1952. Studies on the chromosome number in some Cactaceae species. Ed. by
E.B. Kurtz, Jr. Jap. J. Breed. 1:233-236.

LEUENBERGER, B.E. 1976. Die Pollenmorphologie der Cactaceae. Diss. Bot. 31.

Lona, R.W. and O. Laketa. 1971. A flora of Tropical Florida. Univ. Miami Press, Miami.

NEGRON-OnrtIz, V. 1998. ie ewe biology of a rare cactus, ada sees ica

in the Florida Keys: why is seed set very low? Sexual Pl. Reprod. 11:208-2

Pater, D.T. 1984. Endangered and threatened wildlife and plants; Final a to deter-

mine Cereus robinii (Key Tree-Cactus) to be an endangered species. Fed. Reg. 49(140):29234—

ay,

PEMBERTON, R.W. 1995. Cactoblasti t (Lepidoptera: Pyralidae) in the United States:
An immigrant biological control agent or an introduction of the nursery industry? Amer.
Entomol. 41:230—232.

Pierce, R.L. 1995. Infestation of Opuntia by the phycitid moths Melitara prodenialis and
Cactoblastis cactorum. M.S. Thesis, Florida Atlantic University.

ge D,J., M.A, Baker, B.D. Parrirr, M.W. MOHLENBROCK, and R.D. WORTHINGTON.

985. Chromosome numbers in some Cacti of western North America—V. Syst. Bot.
to: 471-483
7 a - 1930. Consolea corallicola, Florida Semaphore Cactus. Addisonia 15:25—26,

ae NCER, ] L. 1955. A cytological study of the Cactaceae of Puerto Rico. Bot. Gaz. 117:33-

ene, R.P. 1998. Guide to the vascular plants of Florida. University Presses of Florida,
Gainesville

Yuasa, H., H. Suimizu, S. Kasurwai, and N. Konbo. 1973. Chromosome numbers and
their bearing on the geographic distribution of the subfamily Opuntioideae (Cactaceae).
Rep. Inst. Breed. Res., Tokyo Univ. Agric. 4:1-10.

ARKANSAS CAREX (CYPERACEAE): A BRIEFLY
ANNOTATED LIST

FHILIP E avArT

Kisatchie National Forest
2500 Shreveport Highway
Pineville, LA 71360, U.S.A.

ABSTRACT

R llecti d herbarium specimen study resulted in this review of the status of
the 121 taxa in the genus Carex occurring the state of Arkansas, USA. A list of these taxa
provides frequency and habitat data, while county dot maps show known distribution.
The list gives additional information on selected taxa, especially on rare species, on taxa
previously considered rare in Arkansas, or on nine taxa reported new to Arkansas. This
paper considers several taxa as distinct which were previously treated as synonyms by other
authors. It reports four hybrids, and treats taxa excluded from the Arkansas flora, as well
as taxa considered to be possible additions to the state flora.

RESUMEN

tudios d imenes de herbario dieron como resultado esta
revision del estatus 4e los 121 taxa ae género Carex del estado de Arkansas, ce he
lista se tos ue ofrece ones a pecuenels y habitat, ce aus lost mapa d

muestra epccaimes te

de especies raras, de taxa ee raros previamente en es o de fost nueve taxa
citados como anes para Arkansas. Este trabajo considera como distintos varios taxa que
habfan sido tratados como sindénimos por otros autores. Se citan cuatro hibridos, y se tratan
taxa excluidos a la flora de Arkansas, asf como taxa considerados como posibles adiciones
a la flora del estado.

INTRODUCTION

Smith published county record dot maps for 2,469 taxa of vascular plants
(1988), and keys to Arkansas’ vascular flora (1994). In May 1991, the au-
thor set a goal of publishing a book on Arkansas Carex in 2011 and began
field work toward that end. This paper presents initial results of that study
through the 1996 field season. The author's collections, a review of litera-
ture, and a review of collections at several herbaria provide the bases for
this study.

METHODS

Field work in the last seven years resulted in 1523 Arkansas Carex col-
lections, excluding duplicates. 1794 herbarium collections were reviewed

[ACPR had 178 collections, CLEM 18, LSU 24, LSUS 3, LTU 46, MO
Sipa 18(2): 535-554. 1998

536 Sipa 18(2)

149*, MOR 8, NA 88, NLU 652, NO 28, SFRP 21, STAR 147, UAM
113, UARK 237*, UNCC 0, USCH 14, WCUH 1, Buffalo National River
herbarium 58, Sylamore Ranger District/Experimental Forest herbarium
(Ozark National Forest) 9]. An asterisk (*) indicates incomplete reviews.
Anthony A. Reznicek reviewed and provided reports of 483 additional Arkansas

warex collections at MICH, excluding Hyatt collections. Several factors make
the declaration of state records difficult. Consultation with Reznicek re-
sulted in the recognition of many taxa not formerly recognized as distinct
by some authors. Some species pairs previously lumped and now split are
not treated herein as state records. Declaring state records draws attention
to both species new to Arkansas and to species with vouchers which may
have been lying unreported in herbaria.

ECOLOGICAL UNITS OF ARKANSAS

Keys, et al. (1995) provides the basis for a map of the ecological regions
of Arkansas (Fig. 1). Keys’ map and accompanying booklet describe eco-
logical units for the eastern United States, defined as part of the Humid
Temperate Domain. Within Arkansas, the Subtropical and Hot Continental
Divisions of the Domain occur, with the Ozark Highlands and Boston Mountain
Sections represented in the latter, and several sections represented in the
former Division. A brief description of the ecologically defined Sections
represented in Arkansas, and mapped in Figure 1, based, in part, on Keys
text follows. The figure also shows Crowley's Ridge, a significant subsec-
tion. Two letter abbreviations used in Figure 1 precede each section’s de-
scription,
AV—The Arkansas Valley Section contains the Arkansas River in western
Arkansas, and unique Mount Magazine, the highest peak in the state. The
sandy and alluvial soils of the valley contrast with the sandstone and shale
of the Mountain. White Oak, Northern Red Oak, and hickories dominate
the hills which hide a few small wetlands and many perennial streams.

BM—The Boston Mountains, made of high hills and low mountains, lay
over sandstones. Southern Red, White, Northern Red, and Post Oaks share
the hillsides with various hickories, the hills cut by perennial streams.

CP—Often referred to as the West Gulf Coastal Plain, the Middle Coastal
Plains, Western Section (as labeled by Keys et. al. 1995), forms smooth to
irregular plains and riverine valleys on various mixtures of sands, silts, clays,
in gravels. Shortleaf and Loblolly pines mix with a variety of oaks and
hickories, with other wetland species along the frequent streams and rivers.

CR—The subsection known as Crowley’s Ridge, an isolated low ridge of
Pleistocene loess, nearly bisects the Mississipp! Alluvial Basin. Various oaks
as well as Beech-Maple forest thrive on this narrow strip of rolling hills surrounded
by flatlands. Short stream channels drain the ridge.

Hyatt, Arkansas Carex 537

Fic. 1. Ecological units of Arkansas (modified from U.S.D.A. Forest Service, 1995).

MB—The fertile farms of the Mississippi Alluvial Basin Section lie in an
area which once held much of Arkansas’ wetlands. Earthquakes infrequently
rock northeastern sections near the New Madrid fault, causing liquefac-
tion of the alluvial sands, silts, and clays which cover this section. Forests
vary from more upland species such as oaks and hickories, to those associ-
ated with wetter sites to riverine wetlands, including Loblolly Pine, Sweetgum,
hickories, Willow and other oaks, Sycamore, and Birch. Drainage ditches
reduced wetlands, although many wetlands remain.

OU—The Ouachita Mountains, unlike the eroded plain of the Ozarks,
fold into open high hills and low mountains, on sandstone, shales, and cherts,
often with bouldery characteristics. Shortleaf Pine and a mixture of the above
oak species cover the hills, with streams often following the east-west lie of
the mountains.

OZ—The low hills, irregular plains, karst, and entrenched valleys of the
Ozark Highlands contain cherty clay residuum on dolomites, limestones,

538 SIDA 18(2)

and sandstones. Vegetation includes various Oak forests, including White,
Post, Blackjack, and Black Oak, with prairie potential in Bluestem and other
grasses. Entrenched streams and cool springs provide a haven for northern
carices reaching the southern edge of their range.
ARKANSAS CAREX (CYPERACEAE): A BRIEFLY ANNOTATED LIST

‘The following alphabetical list provides a snapshot of current knowledge
on the 121 Carex taxa recognized in Arkansas. Taxonomy follows the draft
of the Flora of North America Carex treatment, provided by A.A. Reznicek
of MICH (ined.). The general format for entries follows:

Accepted genus species (in bold); author(s); figure number (distribution
mapped); syronyms (in italics); frequency, habitat, comments. An asterisk
indicates new state records; voucher specimens cited in some cases.

Frequency statements, unless qualified, represent observed frequency in
Arkansas. Arkansas habitats listed include data drawn from personal ob-
servations, herbarium labels, consultation with others, or in rare cases, the
literature,

The author's collections skew the distributions of some species; Baxter
County, the author’s home for several years, is both the most heavily col-
lected and has the most known species. North Central Arkansas is better
represented than most other regions. Recent collections in south Arkansas,
nearest the author's current home, make it better represented than eastern
Arkansas, a neglected area. For example, Carex umbellata turned up in 28
new western Arkansas counties on a single trip in 1996, while eastern Ar-
kansas has known records primarily om the author’s recent collections.

Arkansas distribution maps have two shades, solid for counties where
the author confirmed the identification of a specimen, cross-hatched when
reported elsewhere without the author's confirmation of the identification.
Most of the identifications of the author’s collections were confirmed or
corrected by A.A. Reznicek, while most herbarium specimens (with the exception
of those at MICH) were not.

—

scondita Mack.—(Fig. 2). Locally numerous, in dense colonies; restricted to flood-

pais in moist deciduous woods (prefers natural levees). See Naczi and Bryson (1990).
*C. aggregata Mack.—(Fig. 2). Rare, moist opening in a disturbed bottom forest,
iad Bice an adj; acent te (formerly forest) and hedgerow by road next to the pasture
(S. e G. Jones, A. & S. Reznicek 11140 vo H, MICH, TRIE, VDB); Soe Co. Bailey
e Gibbons s.n. (WARK) from May 28, 1975 was determined as C. ageregata by Stanley D.
Jones (E.B. Smith, pers. comm.).
*C, alata Torr. & A. Gray.—(Fig. 2). A record of this plant (D. Castaner 9119 at WIS)
from Crowley’s Ridge State Park area, collected May 10, 1986 was pointed out by An-
thony A. Reznicek. This plant, probably scattered in northern Arkansas, also occurs in
Missouri within four miles of Fulton County, Arkansas.
C. albicans Willd. var. albicans.—(Fig. 2). Carex artitecta Mack., C. emmonsii Dew. ex

Torr. var. ay) nbergti (A. Gray) Rettig, C. physorhyncha Liebm. Occasional, widespread;

bluffs, hillsides.

Hyatt, Arkansas Carex

Bi eee beet ne, es

C. alata

\ L
an
|
|
l

be ff 5
C. albicans var. australis

R22 | eo
Tyros 4

i eae | I 5 cial eee Keene
C. albursina C amphibola

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t .
ny >
os ae y, - a
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ot onl ; }
i \ ies fh 4
EY Prise
Tt d } LM

} uA

c. arkansana-

i ieee oes
C. atlantica spp.

a Vs aay
an L—7 SL y eer i
> CLL Pey 4s

a2

C. atlantica sop. capillacea C. austrina

Fic. 2. Documented county records in Arkansas for Carex species (solid = counties where
the author confirmed the identification, and cross-hatched = when reported elsewhere without

the author’s confirmation).

540 Sipa 18(2)

C. albicans Willd. var. australis (L.H. Bailey) Rettig —(Fig. 2). Carex emmonsii Dew. ex
‘Torrey var. australis (L.H. Bailey) Rettig, C. physorhyncha Liebm. Infrequent, poorly known
in state. See Orzell and Bridges (1987).

C. albolutescens is i —(Fig. 2), Occasional to common south, infrequent northwest;
floodplains, wet w

C. albursina Sheldon. —(Fig. 2). Taxonomy of Arkansas material needs study; locally abundant,
deep valleys, mesic north-facing slopes.

C. amphibola Steud. sensu lato—(Fig. 2). Common northwest, occasional elsewhere; val-

leys, floodplains.

C. annectens Bickn. var. annectens.—(Fig. 2). Common (several more counties repre-
eee at UARK per E.B. Smith, pers. comm. 1994), ruderal.

C. arkansana L.H. Bailey—(Fig. 2). Infrequent to rare; prairies, surrounding ruderal ar-
eas (Hyatt 4623.03, 4829.03 MICH, ca

C. atlantica L.H. Bailey subsp. atlantica.—(Fig. 2). Infrequent; streamsides in bog-like
areas, sometimes with Sphagnum mosses. Poorly studied in Arkansas (Orzell 1783 MICH).
C. atlantica L.H. Bailey subsp. capillacea (L.H. Bailey) Reznicek.—(Fig. 2). Locally abundant;
bog-like streamside floodplain (Orzell 1321 MICH).

C. austrina (Small) Mack.—(Fig. 2), Occasional to common north, infrequent south; roadsides,

drairies

"C. basinntha Schl —(Fig. 2), Occasional, locally abundant; floodplains, forms large colonies
along larger streams. Infrequent in the Ozarks, where C. jamesii largely replaces i it (Orzell
1733 MICH). remain distinct from C. willdenowiti. Naczi et al. (1998) discussed the
recognition of this speci

C. bicknellii Britt. var. opaca F.J. Herm.—(Fig. 3). Infrequent; prairie regions and rem-
nants. See Jones and eel (1991) fora key to Texas Ovales, and Herman (1972

C. blanda Dew.—(Fig. 3). Carex laxiflora Lam. var. blanda (Dew.) Boott. Common to
abundant, widespread; mesic woodlands, ruderal.

C. brevior (Dew.) Mack. ex Lunell.—(Fig. 3). Occasional; ruderal, natural habitats poorly
known in Arkansas

C. bromoides Sahie var. bromoides.—(Fig. 3). Rare, cherty soil, full shade (Rettig 1982);
probably from Swany: areas

(Fig. 3): Infrequent to occasional, poorly known; “mesic sandy ravine
pet with some Sites influence” (Orzell & Bridges 1987), floodplain forests.

C. bullata Schk.—(Fig. 3). Rare; wet areas on sandy soil, roadside, open or wooded. Known
in Arkansas from two collections (Moore 400396, and Bryson 5775; UARK). These repre-
sent the only known sites west of the Mississippi River (A.A. Reznicek, pers. comm.). Lipscomb
1980) cites Moore 480070 WIS, from L ogan County.

C. bushiit Mack.—(Fig. 3). Occasional, he ally common; prairies, glades

C. careyana lorr.—(Fig. 3). Infrequent, very locally abundant; deep mesic owalley on talus
slopes often below bluffs (Hyatt 5897 MICH).

C. caroliniana Schwein.—(Fig. 3). Occasional north, common south; ditches, sunny valleys.
C. cephalophora Muhl. ex Willd.—(Fig. 3). Common, widespread; woodlands

C clean Schwein.—(Fig. 3). Common, widespread; ene often of smaller
streams, ditches, old fields, wet woods, swamps.

C. communis L.H. Bailey.—(Fig. 3). Occasional, locally abundant; blufflines, especially
below bluffs on upper talus slopes.

C. comosa Boott.—(Fig. 3). ineecacae borders of larger streams, small lakes.

C. complanata Torr. & Hook.—(Fig. 3). Carex complanata Torr. and Hook. var. complanata
paints north, common to abundant elsewhere; ditches, pine woods, often ruderal.
C.c

onjuncta Boott.—(Fig. 3). Rare; open wooded pasture, wet area (Reznicek 9792 MICH,

—

Hyatt, Arkansas Carex 541

t
|
{

at
g
AA
at

ae E
:

ae

{ Y
Le | t
C. careyana

ae 1 ae

Ee a = =| - at Mba rae
C. comosa C. complanata C. coniuncta

Fic. 3. Documented county records in Arkansas for Carex species (solid = counties where
the author confirmed the identification, and cross-hatched = when reported elsewhere without
the author's confirmation).

Sipa 18(2)

C crebriflora
c davisii

C. crawei
C. crus-corvi

C. corruqata

542

C. decomposita

We
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he mens via

C. debilis var. debilis

i

C, eburnea

igitalis var. macropoda

. dit

C. digitalis var. diaitalis

= counties where

solid
when reported elsewhere

C, fissa var, fissa

(

and cross-hatched

a
oO
o
a
2
n
o
a4
oO

Fic. 4. Documented county records in Arkansas for Carex species

the author confirmed the identification,

without the author’s confirmation).

Hyatt, Arkansas Carex 543

UARK). The author has not seen the voucher (Jones & Reznicek 1995).
C. corrugata Fern.—(Fig. 4). Locally common; restricted to floodplains of major rivers,
streams, low woods.
C. crawei Dew.— (Fig. 4). Occasional, very locally abu t; cal seeps, springs,
glades, most frequent in rights-of-way which cross ne
C. crebriflora Wieg.—(Fig. 4). Occasional to common; ioeal to small drainages, stream
floodplains (Demaree 14415 MICH).
C. crinata Lam.—(Fig. 4). Locally common; perennial streambanks, wetlands.
C. crus-corvi Shuttlew. ex Kuntze.—(Fig. 4). Occasional, often scattered; wetlands, rud-
eral wetlands, often roadsides.
C. davisii Schwein. & Torr—(Fig. 4). Locally common, occasional in north Arkansas,
especially on the Salem Plateau; calcareous stream floodplains, pastures, woodlands (Hyatt
1994
C. debilis Michx. var. debilis —(Fig. 4). Common to abundant south; sweet bay and magnolia
swamps, stream terraces, creekbanks.
C. debilis Michx. var. pubera A. Gray.—(Fig. 4). Rare; Morrow’s Bottoms Natural Area
(Sundell and Barnes 9753, UAM).
Gs a Muhl.—(Fig. 4). Infrequent, swamp and lake margins (Moore 490043
UA
C; oe Willd. var. digitalis —(Fig. 4). Common; upland and north-facing deciduous
woods.
C. digitalis Willd. var. macropoda Fern.—(Fig. 4). Infrequent; rich woods and eas
(Hyatt, et. al. 5290, MICH, UARK). Anthony A. Reznicek reports (pers. comm.) a spec
men collected by Hasse in Little Rock, May 1886, i youn at BH, MICH, and NY.
C. eburnea Boott.—(Fig. 4). Occasional; very local h {ominant, calcareous glade
outcrops, bluffs at wet microsites.
C. emoryi Dew.—(Fig. 4). Rare; streambanks ( Walker, s.n. UARK).
C. festucacea Sebi —(Fig. 4). Common; prairie wetlands, ruderal. Underrepresented in
collections.
*C. fissa Mack. var. fissa.—(Fig. 4). Rare, “Moist, open grassy and sedgey meadow, heavy
clay soil (Reznicek 9271, MICH). See Jones, et. al. (1990).
(Fig. 5). Common; floodplains, widespread in flatlands.
C. frankii i (Eig. 5). Abundant; a ea lakeshores, ditches. Arkansas’ most
widely reported Carex sedge.
C. gigantea Rudge.—(Fig. 5). Infrequent; riverside (Demaree 3379, UARK), lowlands near
Crowley’s Ridge and common in a backwater swamp (Orzell 3483 MICH).
C. glaucescens Ell.—(Fig. 5). Infrequent; floodplains of smaller streams, swamps (A.A.
Martin s.n. SFR
C. glaucodea Tuckerm.—(Fig. 5). Occasional; ruderal, woodlands, frequent in seldom used
woods roads
C. gracilescens Steud.—(Fig. 5). Rare; steep rocky slopes above the Cossatot River (Naczi
1917, 1926, MICH). Record courtesy of A.A. Reznicek.
C. gracillima Schwein.—(Fig. 5). Rare, two very small, local, Ozark populations on moist
north facing rock outcrop (Hyatt 1993), mesic deep valley below sandstone bluff at drip-
ping waterfall (Hyatt 5723, MICH, UARK).
A granularis Muhl.—(Fig. 5). Locally common to occasional; ag seeps, ditches.
C. gravida L.H. Bailey var. gravida.—(Fig. 5). Infrequent; prai
C. grayi Carey.—(Fig. 5). Carex grayii Carey, an orthographic variant. COtenone wooded
wetlands, ditches.
C. grisea Wahl.—(Fig. 5). Occasional; calcareous floodplain woods, less frequent in sur-

C. flaccosperma

aye a = a ee
f 5B uy ©

zi Oe
a } ~} J
if iI\/

wf Ny ED ee oe
vy ,

i,

.

4 ~~

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pt i fa } j
yoy

Cc. ——

C. granularis

~~)
\
{

¢ Ri [
C. hitchcockiana Cc. hyalina

C. hirtitolia
Fic. 5. Documented county records in Arkansas for Carex species (solid = counties where
author confirmed the identification, and cross-hatched = when reported elsewhere

—

the ;
without the author’s confirmation

ToL. S.

Hyatt, Arkansas Carex

C. interior

C. hystercina

C. hyalinolepis

C. jamesii

a
<
D
S
Q
i=
3
=
&
(3)

axiculmis var. copulata

C. latebracteata

C. laevivaginata _

C. laxiflora var. laxiflora v

C. laxiculmis var. laxiculmis

C. lonaii

C. leptalea

C. leavenworthii

Fic. 6. Documented county records in Arkansas for Carex species (solid = counties where

the author confirmed the identification, and cross-hatched

when reported elsewhere

onfirmation).

Cc

without the author's

546 SIDA 18(2)

rounding uplands.
C. hirsutella Mack.—(Fig. 5). Carex complanata Torr. and Hook. var. hirsutella (L.H. Bailey)
a Abundant to common north, occasional to infrequent south; ruderal, very wide-
spre

C " hirifolia Mack.—(Fig. 5). Infrequent; mesic valleys and floodplains of small to me-
iam sized streams at many, but localized sites (Thomas et. al. 1991).

C. hitchcockiana Dew.—(Fig. 5). Infrequent; mesic, topographically shaded and protected,

eys near larger intermittent streams. Such oo streams

—

cool north facing slopes in deep val

often have long — dry sections aa flow under gravel.
C. hyalina Boott.—(Fig. 5). Locally abundant to common absentoutside specialized habitat;
prefers shaded deciduous woods often on clay soils, periodically flooded, usually along

major rivers and tributaries. Often easily found by looking near bridges on rights-of-way
crossing floodplains. See Bryson, et. al. (1 992), Regn el and Naczi (1993), B epnicelk: and
ae (1

C. hyalinolepis Steud.—(Fig. 6). oe forming huge colonies with long, stout rhi-
zomes in eine ditches, wet prairie
C. hystericina Muhl. ex Willd.—(Fig. 6). C aver ell Mubhl. ex Willd. an orthographic
variant. ea very locally abundant; | seeps, springs, and streams
Prefers sunny sites with perennial streams; absent nearly everywhere lees rare at heavily

=
=]

shaded sites.

C. interior . H. Bailey.

and Ball 1980)

C. intumescens Rudge.—(Fig. 6). Common; widespread in wetland woods, wetland edges.

C. jamesii Schwein.—(Fig. 6). Locally abundant; floodplains, especially in deep, narrow

valleys.

C. joorti L.H. Bailey.—(Fig. 6). Common to infrequent; wetland woods, swamps, wooded

roadsides.

C. laevivaginata (Kukenth.) Mack.—(Fig. 6). Infrequent; swampy meadows, calcareous
eeps

(Fig. 6). Rare; calcareous seeps (Orzell and Bridges 1987; Reznicek

C. ibe acteata Waterfall.—(Fig. 6). Infrequent, locally numerous; “well drained dry to
dry-mesic slopes” (Orzell & Bridges, | 87
C. laxiculmis Schwein. var. copulata (L.H. Bailey) Mack.—(Fig. 6). Infrequent; mesic north-
facing slopes | in deep valleys

axiculmis Schwein. var. cena —(Fig. 6). Taxonomy of Arkansas varieties needs

—

study, poorly known, but 1997 collections revealed several new sites; infrequent; mesic
valleys in deciduous woods
: i am. var. laxiflora.—(Fig. 6). Arkansas eueieune. uae understood, Ar-

~p
_

(but vat ; mesic woodlands.

kansas fees needs taxonomic study; occasiona |
2. laxiflora Lam. var. serrulata Underwood.—(Fig. 6). eens distribution poorly un-

ee Arkansas material needs taxonomic study; rare; rich woods.

C. leavenworthii Dew.—(Fig. 6). Common; ruderal.

C. leptalea Wahl.—(Fig. 6). Carex leptalea Wahl. var. leptalea. Occasional; calcareous seeps

in northern Arkansas, wooded seeps at base of slopes in southern Arkansas

C. longii Mack.—(Fig. 6). Occasional oe probably very undercollected, especially in south
Arkansas; wet pais See et east ae 91).
C. louisianica L.H. Bailey.—(Fig. 7). ea swamps and low wetland woods. See
Reznicek and ne (1 1974): \; lexies and Hatch (1990).

ipuliformis Sartwell.—(Fig. 7). Occasional; wetter habitats than and similar habitats
to C lupulina (A.A. Reznicek, pers. comm. 1993).

C. lupulina Muhl.—(Fig. 7). Common; wetland woods, prairie edges, swamps, ditches.

Hyatt, Arkansas Carex 547

C. lurida Wahl.—(Fig. 7). Common; widespread along streams, ae ditches, swamps.
C. meadii Dew.— (Fig. 7). Occasional; limestone glades, sinks, prairi

*C. mesochorea Muhl. ex Willd —(Fig. 7). Reported here as a state Pel for Arkansas;
rare, ruderal; (Hyatt 5485, 5494, 5636. MICH, UARK).

C. microd Torr. & Hook.—(Fig. 7). Infrequent; chalk glades (Moore 400161] UARK),
prairie ee glades. pee Lipscomb (1 980).
C. molesta Mack.—(Fig. 7). Occasional; floodplains, usually of larger streams “commonly
ruderal . .. near or along streams and in open woodland. However it often prefers heavy,
clayey soils and is definitely more of a calciphile [than C. molestiformis]” (Reznicek & Rothrock
1997

=

C. molestiformis Reznicek & Rothrock.—(Fig. 7). Occasional; riverine floodplains, rud-
eral. Reznicek and Rothrock (1997), in recently describing this species, note it “thrives in
a wide variety of open to somewhat shaded, mostly ruderal, habitats with a wide range of
associates, including ditches, dryish roadsides, river and streambanks, moist meadows, and
nf . Typically, colonies of C. molestiformis rarely occur far from streams .
collaborating the author's initial observations. C. molesta, while also exhibiting strong ruderal
tendencies, occurs more typically in sunny upland spring and stream sites aad ruderal set-
tings which imitate these habitats, while C. molestiformis is more riverine.

c. muhlenbergii Willd. var. enervis Boott.—(Fig. 7). Varieties poorly studied in Arkan-
sas; common species; dry, acid soil noclandl late complex (Orzell & Bridges 1987).
muhlenbergii Willd. var. muhlen —(Fig. 7). Varieties poorly eda in Arkan-

sas; common species; widespread in wondiands, ‘roadsides. The varieties may have once
been separate species with separate habitats and little intergrading until recent disturbance
confused the issue (S. D. Jones, pers. c

a meant Schw.—(Fig. 7). Infrequenc floodplains. Orzell and Bridges (1987)
report it in “open areas of a wet-mesic interstream flatwoods.

. nigromarginata oe. —(Fig. 7). including the poorly differentiated C. floridana Schw
Occasional; widespread in woodlands. This species presents a variety of overlapping growth
and color forms, with occasional long rhizomes and pale to heavily purple splotched and
dark ae leaves. Morphology intergrades, making distinctions difficult.

. malis Mack.—(Fig. 7). Reported here new to Arkansas. Rare; seepy roadside ditch

on acid sandstone, deep valley, ee woods, floodplain of the Little Red River of the

Ozark region (Hyatt, et. al. 5275, MICH, UARK), streamside in Sharp County.

C. oklahomensis Mack.—(Fig. 7). i ommon; ditches, wetlands.

C. oligocarpa Schk. —(Fig. 8). Common: streambanks nort th- facing slopes, dec iduou woods.

ouachitana Kral, Manhart, & Bryson.—(Fig. 8). Occasional, locally widespread and

abundant; “arenaceous oak-hickory-pine uplands” (Kral et al. 1987) and “rocky ridgetop

and slope forest throughout much of the Ouachita Mountains” (Orzell and Bridges 1987).

See also McNeilus (1992) and Naczi and Bryson (1990).

C. oxylepis Torr. & Hook. var. oxylepis.—(Fig. 8). Occasional; rich hardwoods in moun-

tains, swamps and ditches elsewhere.

C. oxylepis Torr. & Hook. var. pubescens Underwood.—(Fig. 8). Occasional, locally common;

seeps, moist bluffs, floodplains. Rarely reported outside Arkansas.

C. ozarkana Rothrock & Reznicek.—(Fig. 8). Occasional, locally frequent; Rothrock and

ae (1 396) described this species fromm THe a anicns hills, Oklahoma Ouachitas, and
ne Texas location. The author found it at two locations within 30 miles of, and expects it
n, extreme north Louisiana. Rothrock and Reznicek (ibid) call it “a species of early suc-

cession wetlands on mineral soil . ee: H ea of streams and .. . ditches, pond shores,
and wet depressions in ae, pas

C. pellita Willd.—(Fig. 8). pees ee Michx. Rare (McNalty 321 STAR).

>

SibA 18(2)
dy

y
4
y
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SS
S
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Le
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Py } ae
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C mesochorea

ne | i
ie eee

C. molesta

p—t ~\
} he Nears, rT) Le
Tt Trt

ss

Pi , yy
C, muskinqumensis

° y, \
S t |e. §
enervis C, muhlenberaii var. muhlenberaii

Cc. muhlenberaii var,

i aaa.
Y

(Te beet
C. normalis

Fic. 7. Documented county records in Arkansas for Carex species (solid = counties where
the author confirmed the identification, and cross-hatched = when reported elsewhere without

the author’s confirmati

C. oxylepis var. oxvlepis

C. retroflexa

A cicn ae ~Paqn.
Bat Poi] a
RAL Dee,
| ge Ek Si ee tO E? ‘
of BORG ag grr
a bm tr 4 |
j i — ae oe o
Ms | Lf py 2
S © Loe TL A ett 1S
= Fd ae eS. eu iene ie
bey He Q
ie! N 1 at : «
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30 ei?) (oO

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ae)

i wo
1) Fes ig
2 & § |
3 | : Ra
a 8 4% 3
al ££ ees a2]
a ° (8 s
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r

eat

Me

: ‘scoparia

counties where

Fic. 8. Documented county records in Arkansas for Carex species (solid

the author confirmed the identification, and cross

without the author’s confirmation).

hatched = when reported elsewhere

550 SIDA 18(2)

C. pensylvanica Lam. —(Fig. 8). Occasional; benches on north facing slopes in White Oak

woods, habitat poorly reviewed.

C. planostachys Kunze.—(Fig. 8). Rare, locally abundant; chalk glade outcrop complex.

Northeastern extent of range barely extending into southwest Arkansas in a unique habi-

tat with Juniperus virginiana on exposed white chalk outcrop (Davis and Rettig 2840 APCR

earliest collection found, April 1, 1981).

= prasina Wahl.—(Fig. 8). ie a single collection ae 5050 UARK) from May
1928 along War Eagle Creek in northwest Arlancas “Rare in Mississippi and Mis-

souri’ states A.A. Reznicek (pers. comm. ) suggesting “Crowley's Ridge is a likely bet, along

a woodland seep with Liriodendron.’

*C. radiata (Wahl.) Small—(Fig. 8). Carex rosea sensu Mack. Reported here as new to

Arkansas, rare. Hyatt 5678 MICH, UARK, peh from the Buffalo National River at

TION,R1I5W,S4, in moist calcareous woods near Arkansas Highway 14

C. reniformis (L.H. Bailey) Small.—(Fig. 8). Occasional; floodplains of larger streams,

lakeshores.

C. retroflexa Willd.—(Fig. 8). Common; widespread.

C. rosea Schkuhr.—(Fig. 8). Carex convulata sensu Mack. See Webber and Ball (1984) for

taxonomy and synonymy of C. radiata, C. rosea, and C. socialis, common mountains, in-

frequent elsewhere; valleys and aan are

*C. scoparia Schk. aoe IId.—(Fi 8). ae wet, acmibed ineadow seminates by Juncus,

Carex, and diverse we grasses rere it ft umps (/ez77¢ k 9789 MICH).

Numerous authors meee it Weaee flora eet 1988).

*

. seorsa Howe.—(Fig. 8). Rare; “low woods”. Historic record only, no reports in this
century. A.A. Reznicek (pers. comm. 1995) een a specimen collected by Hasse in 1885
at MICH.

C. shortiana Dew.

(Fig. 9). Infrequent; calcareous sunny wetlands and sprin

C. socialis Mohlenbrock and Schwegmann.—(Fig. 9). Occasional; fige! ele ok larger
rivers in several areas, associated with C. hyalina and C. corrugata. Some authors have treatec
this as a synonym of C. rosea; habitat and morphology make it distinct (Webber & Ball
1984).

C. sparganioides Muhl..—(Fig. 9). Infrequent, typically local in small colonies of fewer
than 20 plants, sometimes 3-4, rarely near 100, with colonies widespread but well scat-
tered; calcareous valleys and floodplains.

C. squarrosa L.—(Fig. 9). Occasional, usually scattered individuals; ditches, floodp
woods.

C. striatula Michx.—(Fig. 9). Locally common; rich wooded slopes on sandy soils. See
Naczi and Bryson (1990).
C. stricta Lam.—(Fig. 9). Infrequent; creek gravelbeds.

C. suberecta (Olney) Britt.—(Fig. 9). Infrequent; calciphile at springs, mossy fens, seeps.
C. swanii (Fern.) Mack.—(Fig. 9). Infrequent; sinks, deep i floodplain

C. texensis L.H. Bailey.—(Fig. 9). Occasional; mountain valley

C. torta Boott.—(Fig. 9). Occasional; mountain gravel streams, aus clumps between
rocks.

C. triangularis Boeck.—(Fig. 9). Common, ditches.

C. tribuloides Wahl.—(Fig. 9). Common; widespread. See also Reznicek (1993).

C. typhina Michx.—(Fig. 9). Occasional; oak savannah wetlands, ditches, creek terraces,
and other wetlands.

C. umbellata Schkuhr.—(Fig. 9). including Arkansas reports of C. abdita. Considered Arkansas’
most frequent and widespread sedge (third most wide

T)

=_

ain

—

—

y reported), easily overlooked, up-
land forests, roadsides. In 1992, ranked $1 (6 or fewer known locations in state)

—

vy the

Hyatt, Arkansas Carex 551

Arkansas Natural Heritage Commission; 1996 collections added 28 counties. Now rival-

ing the frequently collected C. frankti and C. vulpinoidea. Recommended as the state Carex
sedge of Arkansas. Fruits often infested by smut. Early blooming species with fruits often
harvested by ants; may be an important spring food fot ants.

C. virescens Muhl. ex Willd—(Fig. 9). Two records: historic Harvey s.n. UARK, “low woods”,
July 1884, county unknown; Rettig 554 UARK, 20MAY1982, Mont. Co., “Red Oak, Sweet
Gum overstory.

C. aulpinoiie Michx.—(Fig. 10). Abundant; ruderal and widespread, a wetland species.

The second most widely reported Arkansas Carex sedge

C. willdenowii Willd. var. willdenowii.—(Fig. 10). ue known only from a single loca-
tion (Iron Springs Recreation Area, Ouachita National Forest) west of the Mississippi River
(A.A. Reznicek, pers. comm.). See Orzell and Bridges (1987) and Naczi and Bryson (1990).
All other Arkansas “C. willdenowii* specimens reviewed should be considered C. basiantha.

HYBRIDS

C. albicans var. albicans x C. nigromarginata punitive hybrid—Determined
by A.A. Reznicek (Hyatt 6907, MICH); from Woodruff County.

C. flaccosperma x C. glaucodea punitive hybrid—Determined by A.A. Reznicek
(Hyatt 6488, MICH); from Saline County.

C. flaccosperma x C. granularis punitive hybrid—Determined by A.A. Reznicek
(Reznicek 9291, MICH); from Saline County.

C. lurida x C. lupulina punitive hybrid—Determined by A.A. Reznicek
(Hyatt 5720, MICH, UARK); from Lonoke County.

POSSIBLE ADDITIONS

The following taxa, listed as “possible additions” by Smith (1988) have
since been confirmed in Arkansas by collections: C. davisii, C. gracilescens,
C. hyalina (pre-1988 collections of C. hyalina were not represented at UARK),
and C. suberecta. Possible additions to the flora of Arkansas follow.

C. annectens Bickn. var. xanthocarpa (Bicknell) Wiegand.— Hyatt 4343 (MICH, UARK)

was misidentified as this taxon initially. This variety could occur in north Arkansas.
buxbaumi Wahl.—Reported for Arkansas by Demaree (1943). Tucker (1987) notes it

“reaches its southern limit in. . . Arkansas”. Smith (1988) only refers to Tucker (1987)

and lists it as a possible addition to the flora of Arkansas (1994). No vouchers found.

C. incomptera Bickn.—Reported for Arkansas by Demaree (1943). No vouchers found.

C. picta Steud.—Possibly rare in southern Arkansas

C. styloflexa Buckl.—Possibly in “southeast Arkansas in boggy or wet places in hardwoods

at the bases of slopes of bluffs” (Reznicek, pers. comm.).

C. tenera Dew.—Possibly rare in ess pues Unpublished reports by the author

based on an immature specimen of C. ” A

C. verrucosa Muh!.—*“Listed for re by ones (1943); perhaps rare in south Ar-

ine “(Smith 1988).

EXCLUDED TAXA
Smith (1988) excluded the following taxa from the flora of Arkansas; I
concur; C. baileyi, C. douglasii, C. exilis, C. flava, C. gynandra, C. haydenit,

Sipa 18(2)

mA N ea : S \ ~y \ iS ar
Yen i ¢ } — il }
7 To } i : TY Th oat a 2
d aah us bet é {| x

C. shortiana C. socialis C. sparganioides

Lo mt} 4 Te” o a Ie!
RAPA py RAP ALL § py
Ne | | L ; I ey
TAS i | ee 4
‘ey [ iad s | [ A? _

eee t i ite ees [ ff |
. suberecta Cc. swanii C. texensis

Pr,

C. typhina C. umbellata .Virascens

Fic. 9. Documented county records in Arkansas for Carex species (solid = counties where
the author confirmed the identification, and cross-hatched = when reported elsewhere without
the author's confirmation).

Hyatt, Arkansas Carex 553

C. vulpinoidea C. wnlenowe

Fic. 10. Documented county tecondss in gareea for Carex species (solid = counties where
the author confirmed tl ss-hatched = when reported elsewhere without
the author’s confirmation).

C. lacustris, C. muricata, C. plantaginea, C. platyphylla, C. sartwellii, C. sprengelit,
C. tetanica. Two additional taxa, excluded by Smith (ibid), are now known
to occur in Arkansas. Additional excluded taxa follow.

C. backii Boott.—Reported for Arkansas by ee (1980) based on immature speci-

mens of C. latebracteata. A.A. Reznicek (pers. comm.) has seen the voucher.
C. straminea Willd. nny . Soe ee comm. 1996) noted the one Missouri
location for this is a and Oklahoma reports of C. straminea are based

on the recently named C. ozarkana.
C. tetanica Schk.—Anthony A. Reznicek (pers. comm. 1993) notes “not in Arkansas, possibly

not in Missouri even.”

ACKNOWLEDGMENTS

Many thanks to A.A. Reznicek (MICH) for his abundant help in nearly
all facets of the work required to produce this article, E.B. Smith (UARK)
for his continuous encouragement and assistance, and to the various state
and federal agencies who provided permission to collect on their lands, including
especially the U.S.D.A. Forest Service, Buffalo National River, and the Arkansas
State Parks.

REFERENCES

Bryson, C. T.,, R. E C. Naczi, and S$. McDanieL. 1992. Notes on ee records of
Carex (Cyperaceae) from the southeastern United States. Sida 15:1

Demaree, D. 1943. A catalogue of the vascular plants of Arkansas. Taxodium 1:1-88,

HERMANN, EJ. 1972. A new variety of Carex bicknellii from Arkansas. 4

a T, PE. 1993. A survey of the vascular flora of Baxter County, Arkansas. Castanea

15-140.

mae PE. 1994. Significant collections: Arkansas. Castanea 59:78-79.

Jones, S.D, and S.L. Harcu. 1990. Synopsis of Carex Section Lupulinae (Cyperaceae) in
Texas. Sida 14:87-99,

Jones, S.D., G.D. Jones, and J.K. Wiprr. 1990. Carex fissa section Multiflorae (Cyperaceae):
new to Texas. Phytologia 68:47—-50.

Jones, $.D. and A.A. Reznicek. 1991. Carex bicknellii, “Bicknell’s Sedge” ee
new to Texas, with a key to Texas species of section Ovales. Phytologia 70:115-

554 Sipa 18(2)

Jones, $.D. and A.A. REZNICEK. 1995. Carex conjuncta (Cyperaceae) verified for Arkansas,
and notes on the range of Carex oklahomensis. Sida 16:772-774.

Keys, JR., J., C. CarPENTER, S. Hooks, S. KOENIG, F McNas, W.H. Russe, W.E. SMITH,
M-L. 1995. Ecological units of the eastern United States — first approximation (map
and booklet of map unit tables), Atlanta, GA: U.S.D.A. Dept. Agric. Forest Ser. Tech.
Publ. R8-TP 21.

Krat, R., J. MANHART, and C.T. Bryson. 1987. A new Carex sect. Oligocarpae (Cyperaceae)
from western Arkansas and eastern Oklahoma. Ann. Missouri Bot. Gard. 74:440—442

Lirscoms, B.L. 1980. Additions to the Cyperaceae of Arkansas. Castanea 45:70.

McNeEus, V.B. 1992. Carex ouachitana (Cyperaceae) new to Tennessee. Sida 15:154.

Naczi, R.EC. and C.T. Bryson. 1990. Die records of Carex (Cyperaceae) from
the southeastern United States. Bartonia 56:4 .

Naczi, R.ES., A.A. REZNICEK, and B.A. Forp. 1998 Morphological, geographical, and
ecologcal differentiation in Carex willdenowii complex (Cyperaceae). Amer. J. Bot. 85:34—

OrzZELL, S.L. and E.L. Bripces. 1987. Further pars and oo collections in
the flora of Arkansas, with historical, ec ological, a E grat | notes Phytologia
64: 81-144.

RETTIG, J. : 1982. Seven significant vascular plant records from Arkansas. Proc. Ark. Acad.
Sci. 36:9

REZNICEK, i 1990. Evolution in sedges (Carex, Cyperaceae), Canad. J. Bot. 68:1409-
1432

REZNICEK, A.A. 1993. Revision of Carex section Ovales (Cyperaceae) in Mexico. Contr.
Univ. Michigan Herb. 19:97-136.

Reznicek, A.A. and P.W. Ball. 1974. The taxonomy of Carex series Lupulinae in Canada.
Canad. J. Bot. 52:2387-2399.

Reznicek, A.A. and PW. Batt. 1980. The taxonomy of Carex section Stellulatae in North
America north of Mexico. Contr. Univ. Michigan Herb, 14:153-20

Reznicek, A.A. and PE. Hyarr. 1996. Conservation assessment for Carex hyalina. Docu-
ment produced for the U.S. D. A. Forest Service, Kisatchie National Forest, Pineville,
Louisiana.

Reznicrk, A.A. and R.BR.C. Naczi. 1993. Taxonomic status, ecology, and distribution of
mi tag (Cyperaceae). Contr. Univ. Michigan Herb. 19:141-147.

Reznicek, A.A. and P.E. RorHRock. 1997. Carex molestiformis (Cyperaceae), a new spe-
cles o section Ovales from the Ozark Mountain Region. Contr. Univ. Michigan Herb.
21:299—308.

Rorurock, PE. 1991. The identity of Carex albolutescens, C. festucacea, and C. longii (Cyperaceae).
Rhodora 93:51-G66.

Rorurock, PE. and A.A. REZNICEK. 1996. A new species of Carex section Ovales (Cyperaceae)
occurring in the Ozark Mountain region. Brittonia 48:104—110

SMITH, E.B. 1988. An atlas and annotated list of the vascular flora of Arkansas. Published
by the author. Fayetteville, Arkansas (out of print).

SMITH, E.B. 1994. Keys to the flora of Arkansas. University of Arkansas Press. Fayetteville,
Arkansas

Tuomas, R.D., E.B. SmirH, E. SUNDELL, PE. Hyatt, and C. AMason. 1991. Additions to
the flora of Arkansas. Sida 14:483-491.

Tucker, G.C. 1987. The genera of Cyperaceae of the southeastern United States. J. Arnold.
Arbor. 68:361—445.

Webper, J.M. and PW. Batt. 1984. The taxonomy of the Carex rosea group (Section
Phaestoglochin) in Canada. Canad. J. Bot. 62:2058—2073.

AGALINIS (SCROPHULARIACEAE)
IN THE OZARK HIGHLANDS

JOHN EF HAYS

Ozark Environmental Consulting
3308 Keenes Edge Drive
Columbia, MO 65201, U.S.A.

ABSTRACT

The ten species of Agalinis known to occur in the Ozark Highlands are treated floristi-
cally. A key to the species is presented along with descriptions, flowering time, ecological
data, and distribution maps. In addition, taxa considered rare are noted and their current
statuses indicated.

RESUMEN

Se trataron las diez especies de Agalinis conocidas de los “Ozark Highlands.” Se presenta
una clave de las especies, con descripciones, datos fenoldgicos y ecoldgicos, y mapas de
distribucién. Ademas, se anota el estado actual de las especies raras.

INTRODUCTION

Agalinis isa New World genus of about 60 species (Canne-Hilliker 1988).
Thirty-three species of Agalinis occur in North America (Canne-Hilliker
& Dubrule 1993; Canne-Hilliker & Kampny 1991). The genus is most
diversified from southeastern Texas eastward to the Florida panhandle (Canne-
Hilliker & Dubrule 1993; Clewell 1985; Correll & Johnston 1970; McRoberts
1989; Vincent 1982; Wilhelm 1984). Eleven species of Agalinis (sensu Canne-
Hilliker & Kampny 1991) occur in the Midwest (Deam 1940; Holmgren
1986; Mohlenbrock 1986; Smith 1988; Steyermark 1963; Swink & Wilhelm
1994; Voss 1996; Williams 1973), ten of which occur in the Ozark High-
lands. Taxonomically, North American species of Agalinis are relatively well
known, although the need still exists for a modern, monographic treatment
of the genus. Pennell’s work on the group (1929, 1935), while comprehen-
sive and still useful, is also outdated, as recent scholarship has suggested.
Studies by Canne-Hilliker and associates on the cytology, micromorphol-
ogy, comparative anatomy, and seedling morphology of most species of North
American Agalinis (Canne 1979, 1981, 1983, 1984; Canne-Hilliker 1987;
Canne-Hilliker & Kampny 1991; Stewart & Canne-Hilliker 1998) have
resulted in a revision of Pennell’s ideas concerning the delimitations of spe-
cies and the classification of and within the genus.

These revisions, along with nearly 90 years of collecting since Pennell
first undertook his study of the genus, necessitate the presentation of new

SIDA 18(2): 555-577. 1998

556 SIDA 18(2)

data concerning the taxonomic status and distribution of Agalinis throughout
North America. The purpose of my paper is to present an up-to- date ac-
count of the distribution, taxonomy, nomenclature, and rarity of Agalinis
in the Ozark Highlands as well as to provide an unambiguous key for the
identification of these species. Approximately 3,000 herbarium specimens
from 24 herbaria were examined (listed in acknowledgments), and I made
over 200 collections of Agalinis during the course of this study. Each dot
ona distribution map represents one or more verified voucher specimens.
Vouchers I collected were deposited at MO or remain in my possession.

STUDY AREA

The Ozark Highlands, often referred to as the Ozark Plateau(s), Ozark
Province, the Ozark Highland (Cozzens 1940; Fenneman 1938; Sauer 1920;
Thom and Wilson 1980), or locally as simply the * Ozarks,” i is a distinct
pe area classified as the Ozark Plateaus @ province (Thornbury
1965), an area of over 134,000 square once. Within this geomorphic
province fall two distinct ecological sections, the Ozark Highlands Section,
and the Boston Mountains Section (McNab and Evers 1994). The Ozark
Highlands Section comprises over 85 percent of the area, and dolomites
and limestones predominate, ae one of the most extensive karst re-
gions in the country (Nigh et al. 1992). Such extensive karst features as
caves, springs, seeps, and sinkholes are common. In the Boston Mountains
Section to the southwest, on the other hand, sandstones and shales pre-
dominate, and the sandstone exposures that dominate the landscape in this
section stand in contrast to the more carbonate landscape of the Ozark Highlands

Section.

The Ozark Highlands Section is characterized by gently rolling to heavily
dissected hills, whereas the Boston Mountains Section ts characterized by
low mountains, open hills, and hilly plains. The varied topography, as well
as the distinct bedrock and soil types displayed throughout the Ozark Highlands,
has produced a unique assemblage of terrestrial and aquatic communities,
and the Ozark Highlands is considered a center of endemism (Nigh et al.

1992). Broadleaf deciduous forest (mostly oak- hickory) comprises the dominant
vegetation type in the Ozark Highlands, but pine forests, prairies, glades,
fens, marshes, and large rivers are also present in the area and add to its
biological diversity.

The climate of the Ozark Highlands is continental, with precipitation av-
eraging 122 to 132 centimeters annually, and the annual temperature av-
eraging 13 to 18 degrees Celsius (McNab and Evers 1994). Elevation ranges
from a low of 91 meters in Missouri to 914 meters in the Boston Moun-
tains of Arkansas (Keys et al. 1995). Portions of Arkansas, Illinois, Kansas,

Hays, Agalinis in the Ozark Highlands 557
Missouri, and Oklahoma fall within the boundaries of the Ozark High-

lands, but the overwhelming majority of this geomorphic province is found
in Missouri and Arkansas

TAXONOMIC TREATMENT
Agalinis Raf. (nom. conserv.), New Fl. N. Amer. 2:61. 1837. Gerardia L., Sp.
Pie G1 Or 730:

Ours annual, hemi-parasitic herbs with erect stems; leaves opposite or
subopposite, linear to lanceolate or lance-ovate, sessile, entire, or (in two
species) the leaves sometimes 3-cleft, laciniate, or with lateral lobes; inflo-
rescences terminating most branches, a raceme or spike, sometimes racemiform;
late to hemispherical, 5-lobed, the lobes sometimes
reduced to minute teeth; corolla gamopetalous, zygomorphic, 5-lobed, the
two upper lobes reflexed-spreading or projecting forward, the three lower
lobes spreading, usually pale pink to light purple, rarely white, the throat
often dotted purple and with two yellow, longitudinal lines; stamens 4,
didynamous; fruit a loculicidal capsule, globose, subglobose, ovoid, obo-
void, or ellipsoid; seeds small, angular, the testa conspicuously reticulate,
yellowish to black.

Nor th American species are icall difficult at the hological
level, and there is a relatively high percentage of misidentifications, espe-
cially in herbaria where specimens have not been examined critically. This
frequent misidentification is a source of concern, considering the fact that
25 of the 33 North American species of Agalinis are tracked by state natu-
ral heritage programs as sensitive or rare species (Unpublished data, State
Natural Heritage Programs, 1996 and 1997). The status of each species of
Agatlinis considered rare is indicated following its description. Status is based
on the system of ranking rare taxa developed by the Nature Conservancy
(LaRoe 1995). The Nature Conservancy’s current method of ranking is
summarized as follows (G=global and S=state): G1 and $1 (1-5 occurrences
- critically imperiled), G2 and $2 (6-20 occurrences - imperiled), G3 and
S3 (21-100 occurrences - rare), G4 and $4 (more than 101 occurrences -
apparently secure), G5 and S5 (demonstratively secure), GH and SH (of
historical occurrence only), GU and SU (possibly rare, but not enough
documentation exists to propose a ranking), and GX and SX (apparently
extinct or extirpated).

The synonymy listed under each species correlates with the nomencla-
ture of other manuals a worker may use to identify Agalinis in the Ozark
Highlands: Britton & Brown (1913), Fernald (1950), Gleason (1952), Gleason
& Cronquist (1991), Holmgren (1986), Mohlenbrock (1986), and Steyermark
(1963). Nomenclature follows Canne-Hilliker and Kampny (1991).

calyx g,

558 Sipa 18(2)
KEY TO SPECIES

Living plants of Agalinis appear quite different from their pressed and
dried counterparts. It is helpful, therefore, to familiarize oneself with both
fresh and dried material when working with this group. When identifying
fresh or dried specimens, it is often necessary to examine specimens at 10X
or greater magnification, and preferably with a dissecting microscope; sketching
the corolla in the field while fresh to show the orientation of the lobes and
the shape of the tube can also be helpful due to the membranous nature of
the corollas, which can be easily distorted once pressed. The measurements
used in the key and descriptions that follow are based on well-developed
flowering and fruiting specimens, both fresh and dried. One must keep in
mind that species in this genus demonstrate considerable plasticity, and that
immature, depauperate, or diseased plants can make this inherent varia-
tion even more difficult to interpret; to the collector of such specimens:
beware. There is also a genuine need for a thorough and detailed illustra-
tion of each species of Agalinis, which would greatly facilitate proper iden-
tification.

KEY TO THE SPECIES OF AGALINIS IN THE OZARK HIGHLANDS

1. Leaves lanceolate to lance-ovate, 5-25 mm wide, the upper leaves with

two basal, lateral lobes; stem and calyx tube with retrorse-spreading hairs;

calyx 10-16 mm long 2. A. auriculata
. Leaves linear to lanceolate, 0. ac oe 7) mm broad, the upper leaves with-

out basal, lateral lol be glab ionally somewhat

_

occasionally so
scabrous; calyx 3.5-8(—9) mm long.
est pedicels 6 mm or less long, shorter to slightly longer than
the calyx
3. Pedicels 1-3 mm long; calyx lobes 3-6 mm long, cle: o longer
than the calyx tube, the midveins of the lobes prominently ribbed;
leaves lanceolate to linear-lanceolate, lower leaves sometimes 3-
cleft or laciniate . A. heterophylla
3. Pedicels 2-6 mm long; calyx lobes 0.5—2(—2.5) mm long, shorter
than the calyx tube, the midveins of the lobes without prominent
keels; leaves linear to narrowly lanceolate, the lower leaves never
3-cleft or laciniate.
4, Calyx 5-9 mm long, long-campanulate; anthers 1.9-2.5 mm
long; capsule 7-12 mm long, dllipsaid-« to oblong, longer than
broad; leaves narrowly linear, 0.5—1.5 mm wide, erect and as-
cending, harshly scabrous above, the hairs heavily silicified,
oe marbled; fascicled leaves also ascending 1. A. aspera
Calyx 3-5 mm long, hemispheric to short-campanulate; an-
ieee 2.3-3.5 tn mm long; capsule 4-6 mm long, globose to
subglobose, as broad or nearly as broad as long; leaves linear
to linear-lanceolate, 1-4(—5) mm wide, spreading, arching, or
curling, scabrous above, but not harshly so, or the hairs nei-
ther heavily silicified nor appearing marbled; fascicled
spreading, arching, or curling.

oa

eaves

Hays, Agalinis in the Ozark Highlands

559
5. Middle 2/3 of main stem and largest oo noneea a) to
harshly scabrous; b llary fascicles
usually well-developed; 5 side pal veins of the calyx (those
leading to the lobes) as well as the lobes, scabrous ++++++++++ 3. A. fasciculata

5. Middle 2/3 of main eu and largest branches isaztous to
sparingly scabrous, but not harshly
to asce ending, axillary fascicles weakly to well- developed; 5
principal veins of the calyx as well as the lobes usually gla-
brous, or occasionally ve veins of a few capsules scabrous.
6. Middle 2/3 of main stem and largest branches scabrous,
the pubescence obvious, but scattered on both the stem
faces and angles (avoid areas immediately around the nodes);
branches mostly ascending; axillary fascicles usually well-
developed, burs 2 only eee the fascicl ed leaves
=

1
PE g or equaling g length

6. Middle 2/3 of main stem and largest branches glabrous
to ee eS ut pubescence, when present, conspicu -
ous on l around the nodes);
branches mostly spreading, but sometimes ascending, ax-
illary fascicles absent or only weakly developed, the fascicled

leaves, when present, noticeably shorter than the subtending
=

2. Longest pedicels 6-30 mm long, slightly to much longer than the calyx.

7a

if

lyx 5—9 mm long, long- Can ae capsule 7-12 mm long, el-
lipsoid to oblong, longer than broad; pedicels erect -ascending to arched-
ascending; leaves stiffly aire een scabrous above, the hairs

3. A. fasciculata

. A. purpurea

heavily silicified, appearing ma
Calyx 3-5.5 mm long, Tease to campanulate; capsule 3.5—6
(~7) mm long, globose or obovoid, as broad as long; pedicels spreading
to ascending; leaves spreading to ascending, subglabrous to scabrous,
but if scabrous, the hairs neither heavily silicified nor appearing marbled.
8. Upper two lobes of the corolla arching or projecting forward over

the stamens; eoror4 eos aaa OOS in base of the two
“11

Upper lobes; ome b fascicles

often prese

9. Stem Cae leaves narrowly linear, 0.8-1.5 mm wide; the
two upper lobes of the corolla projecting forward over the sta-
mens, half the length of the three lower lobes, 2-3 mm long;
pedicels scabrous at least basally; anthers 3-4 mm long, the
two upper filaments densely villous basally and distally; seeds
0.8—1.6 mm long, the testa cells large in proportion to the total

[

1. A. aspera

size of the seed 6. A. homalantha

bee)

. Stem glabrous or with a few scattered hairs on the angles; leaves
linear to narrowly lanceolate, 1-5(—6) mm wide; the two up-
per lobes of the corolla arched over the stamens, subequal to
the three lower lobes, 4—5 mm long; pedicels glabrous; anthers
1—2.3 mm long, the upper filaments sparingly hairy through-
out, but not densely villous basally and distally; seeds 0.7—1
mm long, the testa cells small in proportion to the total size of

the see 9. A. tenuifolia

560 Sipa 18(2)
8. Upper two lobes of the corolla reflexed-spreading; corolla pubes-
cent within as the base of the two upper lobes; pedicels not
abr up rtly l] y (occasionally a few pedicels of A, gattingert
are upcurved een Bscicleoabsent,
10. Outside of the three lower corolla lobes pubescent; leaves spreading,
arching, or curling; numerous floriferous, lateral branches present,
the ace appearing, solitary at the mpee) a terminal maces

TdlCl

o 4-angled 4. A. gattingeri
10. Ou of the three lower corolla lobes glabrous; leaves mostly
erect and ascending; few to many branches bearing obvious
terminal racemes present; stem noticeably 4-angled and appearing
wingec

a

1. Stem scabrous on the angles; few to many strongly ascending
branches present; calyx hemispherical, the longest lobes 0.7—
1.2 mm long; fruit globose 8. A. skinneriana
11. Stem glabrous on the angles; a few to many widely spread-
ing branches present; ci aly x campanulate, the longest lobes
ic 2.2 mm long; fruit obovoid 10. A. viridis

1. Agalinis aspera (Douglas ex Benth.) Britton. Gerardia aspera Douglas ex Benth.

Stem 2—7 dim, subglabrous to sparingly scabrous, the branches ascending.
Leaves narrowly linear, stiffly ascending, 0.5—1.5 mm wide, 1.5—4 cm long,
harshly scabrous above, presenting a marbled appearance, and frequently
scabrous on the midveins below. Fascicles well-developed and also ascending.
Inflorescences short to elongate racemes. Pedicels erect to arched-ascending,
stout, 5-11 mm long, to 16 mm in fruit. Ca/yx campanulate, 5—9 mm long,
the lobes 1.0-2.5 mm long. Corolla deep pink, 16—25(—28) mm long, the
throat glabrous within or with a few scattered hairs across the base of the
two reflexed-spreading, upper lobes. Anthers 1.9-2.5 mm long. Capsule ellipsoid
to oblong, clearly longer than broad, 7-12 mm long. Seeds black, 0.9-1.1
mm long. 7=14 (Léve & Léve 1982). Note: voucher not seen. Flowering
late August—early October.

Distribution.—Dry prairies, open woodlands, glades, and blufts. Very
rare in the Ozark Highlands (Fig. 1)

Ecology and conservation status.—GS5; $2 in Arkansas. Although this
species has been reported from Arkansas (Smith 1988), I have not found
any vouchers specimens; all specimens labeled A. aspera from Arkansas have
proved to be another species. Agalinis aspera is largely a species of the eastern
Great Plains, and populations found eastward of this area are mostly peripheral
and of scattered occurrence, usually being restricted to xeric habitats, such

as blufts, glades, and loess hills.

2. Agalinis auriculata (Michx.) S.F. Blake. Gerardia auriculata Michx., Orophylla
auriculata (Michx.) Small, Zomanthera auriculata (Michx

—
—

al,

Hays, Agalinis in the Ozark Highlands 561

Stem 2—7(-8) dm, retrorsely scabrous, simple to sparingly branched above.
Leaves lanceolate to lance-ovate, the upper with two basal, lateral lobes, 2.3—
6 cm long, 0.5—2(—2.5) cm wide, scabrous above and below, lateral veins
visible on both surfaces. /nflorescences spikes. Flowers essentially sessile, oc-
curring in pairs in the upper leaf axils. Ca/yx campanulate, 10-16 mm long,
the tube spreading-pubescent, the lobes 6-12 mm long, scabrous, longer
than the tube. Corolla pink, 20—27(—29) mm long, the throat pubescent
within across the base of the two reflexed-spreading, upper lobes. Azthers
1.9-2.4 mm long. Capsule ovoid, 9-14 mm long. Seeds yellowish, 1.2-1.6
mm long. 2=13 (Kondo 1973). Flowering late August-September.

Distribution.—Dry or mesic prairies, fallow fields, borders of dry woods,
and roadsides. Very rare in the Ozark Highlands (Fig. 2).

Ecology and conservation status. —G3, $1 in Arkansas, $2 in Illinois, $2
in Kansas, S2 in Missouri, and $1 in Oklahoma. Although a rare species
throughout its range, Agalinis auriculata apparently can tolerate consider-
able disturbance, as evidenced by its ability to persist in degraded prairies,
fallow fields, grazed meadows, and roadsides. Despite this apparent ablility
to tolerate continued human disturbance, the Jasper, Newton, Ozark, and
St. Louis County, Missouri, sites, as well as the St. Clair County, Illinois,
site, are based on historic collections or the populations are thought to be
extirpated. Fortunately, however, there are a number of recent collections
from the northern half of Missouri.

Ds Agalinis seicalats eon Raf. Gerardia fase iculata Elliott. Gerardia fasciculata
Elliott f. fasciculat.

Gerardia fasciculata Elliott f. albiflora Palmer.

Stem 2.5-12 dm, scabridulous to harshly scabrous, much branched on
larger plants, the branches ascending or spreading. Leaves linear to narrowly
lanceolate, 1-4 cm long, 1—-2(—4) mm wide, spreading, arching, or curl-
ing, scabrous above and often on the midveins below. Fascicles present and
well-developed or sometimes only weakly so, the fascicled leaves shorter
than to nearly approaching the main leaves, also spreading, arching, or curling.
Inflorescences short to elongate racemes. Pedicels 2-6 mm long. Calyx cam-
panulate to hemispherical, 3-5(—6) mm long, the lobes 0.5—1.5(—2) mm
long, the five principal veins of the calyx (leading to the calyx lobes) sca-
brous; corolla pink to light purple, rarely white, 15-30 mm long, the throat
noticeably pubescent within across the base of the two reflexed-spreading,
upper lobes. Anthers 2.3-3.5 mm long. Capsule subglobose, 4.5—6(—7) mm
long. Seeds 0.6-1 mm long, dark brown to black. 7=14 (Canne 1981). Flowering
from late August—late September.

Distribution.—Dry to moist prairies, borders of woods and ponds, open

ground, fallow fields, and roadsides. Scattered in the Ozark Highlands (Fig.

SipA 18(2)

Fic. 1. Presently confirmed county records in the Ozark Highlands for Agalinis aspera.

3), occurring most frequently in the west and southwest portions of our
area, where it nears the northern and northwesternmost limit of its range.

Ecology and conservation status —G5; S2 in Kansas. In our area, A. fasciculata
is most easily confused with the closely related and widespread A. purpurea
(L.) Pennell. Less scabrous specimens of A. fasciculata with weakly devel-
oped fascicles are especially troublesome and difficult to separate from the
more fascicled and scabrous specimens of A. purpurea (fortunately few specimens
pose such problems). It is not the amount of stem pubescence on the less
scabrous specimens of A. fasciculata that best distinguishes it from the more
scabrous specimens of A. purpurea, but rather the size of the hairs, which

Hays, Agalinis in the Ozark Highlands 563

tend to be larger and broader-based, and in the distribution of the pubes-
cence, which is usually scattered, but present throughout the stem faces as
well as on the angles of the main stem in A. fasciculata.

Agalinis purpurea, on the other hand, has smooth stem faces and quite
often smooth stem angles. On a noticeably pubescent plant of A. purpurea,
the broad-based hairs occur mostly along the stem angles at regular inter-
vals. It is important to rely on the middle two-thirds of the main stem and
largest branches when making observations concerning stem pubescence,
as the smaller branches on both species can be quite scabrous, especially
distally. In addition, the nodes should be avoided when relying on pubes-
cence in identification, because both species are usually quite scabrous around
these areas.

4. Agalinis gattingeri (Small) Small ex Britton. Gerardia gattingeri Small.
Agalinis skinneriana (sensu Holmgren 1986).

Stem 1—5 dm, subglabrous to slightly scabrous, with spreading to some-
what ascending branches. Leaves narrowly linear, 0.4-1.4 mm wide, 1.5—
3(—4) cm long, spreading, arching, or curling, slightly scabrous above, and
often on the midveins below. Fascicles absent. Inflorescence racemiform, a
true raceme rarely developed, most flowers appearing solitary at the nodes
on numerous short, floriferous, lateral branches. Pedicels slender, spread-
ing-ascending, 4—25(—30) mm long. Calyx hemispherical, 3-5 mm long,
the lobes 0.5—-1.4 mm long. Corolla pink to light purple, 8-15 mm long,
the throat noticeably pubescent across the base of the two reflexed-spread-
ing, upper lobes, the outside of the three lower lobes pubescent. Anthers
1.3-2.0 mm long. Capsule subglobose, 4-5 mm long. Seeds 0.5—0.9 mm
long, yellowish. m=13 (Canne 1984). Flowering late August—early October.

Distribution.—Dry, open woodlands, often in rocky soils of chert or sandstone,
borders of dry forests, dry roadsides, glades, bluffs, and exposed ridges. Common
throughout the Ozark Highlands (Fig. 4).

Ecology and conservation status.—None. Holmgren (1986) combined A.
gattingert with A. skinneriana, but the two species have been shown to be
distinct based on macromorphological and anatomical evidence (Canne-
Hilliker 1987; Canne-Hilliker and Kampny 1991). Agalinis gattingeri and
A, tenuifolia are the two most common species in the Ozark Highlands and
are the only two species that have established themselves throughout the
region with any frequency. Our other species occur mostly on the periph-
ery of the Highlands or are localized within the area in particular habitat
types.

5. Agalinis heterophylla (Nutt.) Small ex Britton. Gerardia heterophylla Nutt.

Stem 3-8 dm, glabrous to very sparingly scabrous, usually much branched

564 Sipa 18(2)

Fic. 2. Presently confirmed county records in the Ozark Highlands for Agalinis auriculata.

above. Leaves linear-lanceolate to lanceolate, 1.5—3.5 cm long, 1.8-6 mm
broad, the lowermost sometimes 3-cleft or laciniate, but these usually not
present at flowering time, scabrous above and occasionally on the midveins
below, the undersurfaces of larger stem leaves with a few faint, but notice-
able, lateral veins, (these veins usually not noticeable on dried specimens).
Fascicles absent. Inflorescences short to elongate racemes. Pedicels stout, 1-3
mm long. Calyx campanulate, 5.5—10(-11) mm long, the lobes 3-6 mm
long, clearly longer than the calyx tube, usually with a prominent midnerve
and appearing ribbed. Corolla pink to light purple, 20-33 mm long, the

throat glabrous across the base of the two reflexed-spreading, upper lobes.

Hays, Agalinis in the Ozark Highlands 565

Anthers 2.7-3.5 mm long. Capsule subglobose, 5—8(—9) mm long. Seeds dark
brown to black, 0.7—1.1 mm long. m=14 (Canne 1981). Howering late August—
September.

Distribution —Moist prairies, fallow fields, moist, open ground, and roadsides,
usually in moist soils, but occasionally found in drier sites. Rare and local-
ized in the Ozark Highlands (Fig. 5).

Ecology and conservation status. —G5; $2 in Kansas, S1 in Missouri. This
is another species of Agalinis that nears the northwesternmost occurrence
of its range in the Ozark Highlands; it is much more common in southern
Arkansas, eastern Texas, Louisiana, and Oklahoma outside of the Ozarkian
region of the state.

6. Agalinis homalantha Pennell. Gerardia homalantha (Pennell) Pennell.

Stem 3-10 dm, scabrous. Leaves narrowly linear, scabrous above and on
the midveins below, 1.5—4.5 cm long, 0.8-1.5 mm wide. Fascicles well-developed.
Inflorescences short racemes, with only 3—8(—10) flowers per raceme. Pedicels
slender, spreading-ascending, often upcurved distally, 6-30 mm long, usually
scabrous throughout, but at least basally. Calyx campanulate, 3—5(—6) mm
long, the lobes 0.6-1.5 mm long. Corolla deep pink, 15-26 mm long, the
throat glabrous across the base of the two flattened to slightly arched, for-
ward projecting upper lobes, the upper two lobes 2—3 mm long and half
the length of the three spreading lower lobes. Anthers 3-4 mm long, the
upper filaments densely pubescent basally and distally. Capsule globose, 5—
7 mm long. Seeds dark brown to nearly black, 0.8-1.6 mm long, 7=13 (Vincent
1982). Flowering August-September.

Distribution.—Dry, sandy soils along roadsides, along the dry, sandy banks
of the Arkansas River and the dry to xeric, sandy terrace communities just
beyond the banks of the river. Very rare in the Ozark Highlands (Fig. 6),
restricted to a few counties where the Ozark Highlands approach the Ar-
kansas River Valley in Conway, Crawford, Franklin, and Pope counties in
Arkansas, and Sequoyah County in Oklahoma. This is yet another species
of Agalinis that reaches the limits of its range in the Ozark Highlands; in
this case the species northernmost occurrence.

Ecology and conservation status.—G5; $1 in Arkansas, $1 in Oklahoma.
This species was only recently found to occur in Arkansas, and most of the
sites in the state are found outside of the Ozark Highlands along the Ar-
kansas River. I believe based on my field work along the Arkansas River
alone that enough sites will eventually be found to judge this species de-
monstratively secure in Arkansas. Agalinis homalantha has also been col-
lected recently in Mississippi. While reviewing the holdings of Agalinis at
the Missouri Botanical Garden in January 1997, I found a specimen origi-
nally determined as A. tenuifolia that is actually A. homalanta; this collec-

566 SIDA 18(2)

} _=—2ee: ee all

ee ee
a

\!
oe
4
we
ie
i

WA
a

i
~

Fic. 3. Presently confirmed county records in the Ozark Highlands for Agalinis fasciculata.

tion represents the first known occurrence for this species in Mississippi.
MISSISSIPPI: Bolivar Co.: 2 mi S of Rosedale, W of MS HW 1, 1T23N,
R8W, S. 27, $2; coarse, sandy soil along wood’s edge between levee and
Mississippi River, 16 Aug 1990, Bryson 10170 (MO).

In our area this species is most easily confused with A. tenuifolia, from
which it can be readily distinguished by its scabrous stems and pedicels,
the two very short, flattened or slightly arched upper corolla lobes, the size
of the anthers, which average twice as long as those of A. tenuifolia, and
the much larger seeds.

Hays, Agalinis in the Ozark Highlands

|

=

[
a
re]

=

|
]
4

ae

—

ands for Agalinis gattingeri.

Fic. 4. Presently confirmed county records in the Ozark High

7. Agalinis purpurea (L.) Pennell. Agalinis purpurea (L.) Pennell var. purpurea. Gerardia
purpurea L. var. purpurea. Gerardia purpurea L. f. purpurea.

Gerardia purpurea L. f. albiflora Britton.

Stem 1-10 dm, glabrous to subglabrous, much branched on larger plants.
Leaves linear to narrowly linear-lanceolate, 1-4(—5) mm wide, 1—5 cm long,
spreading, arching, or curling, scabrous above and often on the midveins
below. Fascicles usually absent or only weakly developed, but when present
clearly shorter than the main leaves. /nflorescences short to elongate racemes.
Pedicels 2-5(—6) mm long. Calyx campanulate to hemispherical, 4-6 mm

—

568 Sipa 18(2)

Lenae beternnbylla
fs

Fic, 5. Presently confirmed county records in the Ozark Highlands for Ag

long, the lobes 0.8—2(—2.3) mm long, the five principal veins of the calyx
(leading to the calyx lobes) glabrous or rarely a few veins slightly scabrous.
Corolla rose-purple, rarely white, 18-33 mm long, the throat noticeably
pubescent within across the base of the two reflexed-spreading, upper lobes.
Anthers 2-3.5 mm long. Capsule subglobose, 4—6 mm long. Seeds dark brown
to black, 0.7—1.2 mm long. n=14 (Canne 1981). Flowering late August—
early October.

Distribution.—Moist prairies and roadsides, moist open ground, ditches,
and pond margins. Rare and scattered in the Ozark Highlands (Fig. 7),

but probably more common than its distribution suggests.

Hays, Agalinis in the Ozark Highlands 569

L peed + eae

Fic. 6. Presently confirmed county records in the Ozark Highlands for Agalinis homalantha.

Ecology and conservation status.—G5; $2 in Kansas, $2 in Missouri. Agalinis
purpurea and A. fasciculata are the most difficult species to distinguish from
one another in the Ozark Highlands due to the variable nature of each taxon
(see the discussion under A. fasciculata). Although A. fasciculata can be found
under wet to mesic conditions, A. purpurea cannot tolerate the more xeric
conditions favored by A. fasciculata in our area, and thus a difficult speci-
men collected on an upland prairie or another dry to xeric site can be de-
faulted to A. fasciculata. Occasionally, A. purpurea is found in dry soil, but
close attention to its associate species will indicate that the habitat is actu-
ally a seasonally dry, moist to wet community.

570 Sipa 18(2)

SOY
a hi
oF
“| |
|
ry
\

Fic. 7. Presently confirmed county records in the Ozark Highlands for Agalinis purpurea.

8. Agalinis skinneriana (A.W. Wood) Britton. Gerardia skinneriana A.W. Wood.

Stem 1—5(—6) dm, sharply 4-angled, appearing winged, the angles sca-
brous but the stem faces glabrous, simple to sparingly branched, the branches
ascending. Leaves linear, ascending-appressed, the lowermost slightly spreading,
0.5—1.5 mm wide, 1—2(—2.5) cm long, scabrous above and on the midveins
below. Fascicles absent. /nflorescences short racemes. Pedicels 0.4—1.6(—2) cm
long. Calyx hemispherical, 2-4(—4.5) mm long, the lobes 0.4—1.2 mm long.
Corolla pink to rarely white, 8-16 mm long, the throat noticeably pubes-
cent within across the base of the two reflexed-spreading, upper lobes, the
outside of the three lower lobes glabrous. Anthers 0.6—1.2 mm long. Cap-

Hays, Agalinis in the Ozark Highlands

Fic. 8. Presently confirmed county records in the Ozark Highlands for Agalinis skinneriana.

sule globose, 3.5—5 mm long. Seeds 0.6-1.0 mm long, yellowish. 7=13 (Canne
1984). Flowering late August—early October.

Distribution.—Dry prairies and large dolomite glades. Rare and local-
ized in the Ozark Highlands (Fig. 8).

Ecology and conservation status. —G3; $2 in Illinois, $3 in Missouri. Al-
though not state-listed in Kansas, there are only a handful of known sites
there, and the species clearly warrants protection in that state. Smith (1988)
reported A. skinneriana from numerous counties in Arkansas, but I have
been able to locate only one historic (19th century) specimen of this spe-
cies for the state in all of my herbarium searches, and the specimen has

572 Stipa 18(2)

very imprecise label data. ARKANSAS: Prairies and woods, N.W. Arkan-
sas, Floyd s.n. (UARK); all other specimens seen by me labeled A. skinneriana
have proved to be another species. The sites in Barry and Iron counties,
Missouri, are based on historic collections, but I believe that A. skinneriana
will eventually be relocated in Iron County or found in adjacent Reynolds
County, due to the abundance of suitable habitat (in this case igneous glades)
tha still remains in both counties. Steyermark (1963) also recorded A. skinneriana
from McDonald County, Missouri. He was undoubtedly referring to a specimen
collected by B.E. Bush, cited by Pennell (1929), that resides in the herbarium
of the Botanical Museum at the University of Copenhagen, Denmark (C).
But because | did not review this specimen I have not included A. skinneriana
from McDonald County, although Pennell’s annotation was in all likeli-
hood correct.

On dry prairies A. skinneriana occurs mostly where the vegetation is sparse,
such as the summit of hills or on gentle slopes where the substrate is near
the surface, or along the upper parts of the narrow draws that frequently
occur on prairies, where some slight erosion is usually evident. On dolo-
mite glades it is found in either the more vegetated parts of the glade or in
nearly bare soil.

In the Ozark Highlands Agalinis skinneriana is often confused with A.
gattingeri, or smaller plants of A. tenuifolia. It is readily distinguished from
A. tenuifolia by the corolla, which in A. skinneriana is pubescent within
across the base of the two reflexed-spreading, upper corolla lobes, and by
the sharply winged stem with its noticeably scabrous margins. It can be
distinguished from A. gattingeri by the presence of a true raceme, the gla-
brous outersurface of the three lower corolla lobes, and erect-ascending leaves.

9. Agalinis tenuifolia (Vahl) Raf. Gerardia tenuifolia Vahl. Gerardia tenuifolia Vahl
var. tenuifolia, Gerardia tenuifolia Vahl subsp. tenuifolia.
Agalinis besseyana (Britton) Brittor
Agalinis tenuifolia (Vahl) Raf. var. ny ae (Benth.) S.F. Blake. Gerardia tenuifolia
Vahl var. macrophylla Benth. Gerardia tenuifolia Vahl subsp. macrophylla (Benth.) Pennell.
Agalinis tenuifolia (Vahl) Raf. var. parviflora (Nutt.) Pennell. Gerardia tenuifolia Vahl
var. parviflora Nutt. Gerardia tenuifolia Vahl subsp. parviflora (Nutt.) Pennell.
Stem 2—7 dm, subglabrous to glabrous, much branched on larger plants.
Leaves linear to lanceolate, spreading, ascending, or arched, 1—5(—6) mm
wide, 3-7 cm long, scabridulous above and often on the midveins below.
Fascicles absent or present. /nflorescences elongate racemes. Pedicels slender,
spreading, often abruptly upcurved distally, 6-25 mm long. Calyx hem1-
spherical, 3-5.5 mm long, the lobes 0.5—2 mm long. Corolla light to dark
pink, 10-16 mm long, the lobes 4-6 mm long, the throat glabrous within
across the base of the two forward arching, upper lobes, the two upper lobes
subequal to the three lower lobes. Azthers 1-2(—2.3) mm long. Capsule globose,

Hays, Agalinis in the Ozark Highlands 573

: | Tl pace oS

Fic. 9. Presently confirmed county records in the Ozark Highlands for Agalinis tenuifolia.

4—6 mm long. Seeds tan to dark brown, 0.7—1 mm long. n=14 (Canne 1984).
Flowering principally from late August—early October, occasionally flower-
ing as early as late June.

Distribution.—Dry or moist prairies, borders of woodlands, ponds, marshes,
and rocky streams, also in ditches, fallow fields, and along railroads and
roadsides Common in the Ozark Highlands (Fig. 9), but less frequent in
the west and southwest portions of the region.

Ecology and conservation status —G5; $3 in Oklahoma. Along with Agalinis
gattingeri, A. tenuifolia is the most common species in the Ozark High-
lands, and is the most common and widespread species of Agalinis in North

574 Stpa 18(2)

America. Agalinis tenuifolia and A. gattingeri are the only two species that
occur with any frequency in the Ozark Highlands, the other species being
restricted to the periphery of our area or to Jahr oa habitats within the
Highlands. Agalinis tenuifolia is easily recognized in the field during flow-
ering by the two long, forward arching, upper lobes of the corolla, which
project over the anthers and most of the style. The only species that it can
be confused with in the field during flowering in our area is A. homalantha.
In A. homalantha, however, the style extends well beyond the two very short,
forward to slightly arching upper corolla lobes, the pedicels are scabrous,
and the species has a specific habitat and restricted range in our area.

Agalinis tenuifolia | is a complex and variable species for which a number
of intergrading varieties have been described. As Steyermark (1963) aptly
stated of the varieties in our area: “he varieties presented here are not very
distinct, except in their extremes, often showing overlapping characters, thus
making their determination doubtful or unsatisfactory.” Although it is possible
at times to identify plants confidently as a particular variety, I believe it
best at present to treat these weak varieties as one highly variable species
pending future research.

10. Agalinis viridis (Small) Pennell Gerardia viridis Small.

Stem 1—6 dm, sharply 4-angled, appearing winged, the angles and stem
faces glabrous, with a few to many widely spreading branches. Leaves lin-
ear 0.5—2 mm wide, sometimes the lowermost stem leaves approaching 3
mm in width, 1—2.5(—3) cm long, scabrous above and on the midveins below.
Fascicles absent. Inflorescences racemes. Calyx campanulate, 3—4.5 mm long,
the longest lobes 1.5—2.2 mm long. Corolla translucent to pink, 8-12 mm
long, the throat noticeably pubescent within across the base of the two re-
flexed-spreading, upper lobes, the outside of the three lower lobes glabrous.
Anthers 0.8-1.3 mm long. Capsule obovoid, 5—6(—7) mm long. Seeds yel-
lowish, 0.7—1 mm long. 2=?. Flowering late August-September.

Distribution.—Moist prairies, along the non-forested portions of prai-
rie draws, along the borders of seasonally wet depressions in prairies where
the vegetation is sparse, and rarely on the drier portions of prairies. Very
rare in the Ozark Highlands (Fig. 10), where it is currently known from
only four counties.

Ecology and conservation status. —G4; $1 in Missouri, $1 in Oklahoma.
Agalinis viridis is yet another species that reaches its northern and
northwesternmost occurrence in the Ozark Highlands; it is much more common
south of the Arkansas River, and occupies a wider range of habitats there.
The Jasper County, Missouri, location represents an isolated occurrence of
this species; it was once collected throughout this county, but is now re-

Hays, Agalinis in the Ozark Highlands

rd T Ve
4 |

pret

ae eee

Fic. 10. Presently confirmed county records in the Ozark Highlands for Agalinis viridis.

stricted to Wah-Sha-She Prairie Natural Area, over 300 kilometers from
the nearest known extant site for this species.

ACKNOWLEDGMENTS

I thank Jim Harlen of the University of Missouri’s geography depart-
ment for his help with distribution maps; Judith Canne-Hilliker of the University
of Guelph in Ontario, Canada, for sharing her fathomless knowledge of
Agalinis with me during our frequent communications; John Thieret of Northern
Kentucky University, Floyd Swink of the Morton Arboretum in Lisle, Ili-
nois, and George Yatskievych of the Flora of Missouri Project at the Mis-

576 SIDA 18(2)

sourl Botanical Garden for critiquing the manuscript; lom Foti and John
Logan of the Arkansas Natural Heritage Commission for pointing me to
areas of interest in Arkansas; William Hess of the Morton Arboretum and
Robin Kennedy of the Dunn-Palmer Herbarium at the University of Mis-
souri-Columbia for procuring loans as well as providing working space; and
the curators of the following herbaria for providing me with loans or al-
lowing me to visit their herbaria: APCR, DSC, FE ILL, ILLS, KANU, KSP,
MISS, MO, MODNR, MOR, NLU, OKL, SEMO, SIU, SOTO, STAR,
SMS, TULS, UAM, UARK, UCAC, UMO, and WARM.

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—-. 1982 a taxonomic significance of seedling morphology in Agalinis (Scrophulariaceae).
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—--. 1984. C Seaeeae numbers and the taxonomy of North American Agalinis
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—

578 Sipa 18(2)
BOOK NOTICE

JOHN SHELHAS and RussELL GREENBERG, eds. 1996. Forest Patches in Tropical
Landscapes. ISBN 1-55963-426, pbk; ISBN 1-55963-425-1, hbk).
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promote forest patches as a mechanism to conserve
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—_—

well. | highly recommend this book wit

SIDA 18(2): 578. 1998

FLORA ORNAMENTAL EN PLAZAS Y JARDINES
PUBLICOS DEL AREA METROPOLITANA
DE MONTERREY, MEXICO

ALEJANDRA ROCHA ESTRADA, TERESA E. TORRES CEPEDA,
MA. DEL CONSUELO GONZALEZ DE LA ROSA, SALOMON J.
MARTINEZ LOZANO y MARCO A. ALVARADO VAZQUEZ
Departamento de Botdnica
Facultad de Ciencias Bioldgicas
Universidad Auténoma de Nuevo Leon

Apdo. Post. 2790, Monterrey, N.L., MEXICO

RESUMEN

Se realizé un inventario de las especies vegetales que son utilizadas con fines ornamentales
en el drea metropolitana de Monterrey, encontrandose que se cuenta con 137 especies
pertenecientes a 68 familias, de las cuales Fraxinus sp., Ligustrum lucidum Ait., Rosa centifolia
L. y Melia azedarach L. son las especies mds frecuentes.

ABSTRACT
An inventory was taken of the ornamental eee in the metropolitan area of Monterrey.

We found 137 species in 68 families. The most frequent species are Fraxinus sp., Ligustrum

lucidum Ait., Rosa centifolia L. and Melia azedarach

INTRODUCCION

Es indudable la importancia ecoldégica y estética de las plantas ornamentales
en las ciudades modernas, donde se conjuntan infinidad de problemas que
son el resultado de la urbanizacién exhaustiva, y en las cuales las plantas
ounament ales pasan en el mor de los casos a segundo término, sin considerar
los mul iples b tan, como son: purificaci6n de la atmésfera

Iecvalescontaminada: por el mismo hombre; liberacién de oxigeno; evitan
la erosidn del suelo, pueden funcionar como cortina rompeviento, etc. Desde
el punto de vista estético, las plantas aumentan la belleza de la ciudad con
su follaje multicolor, hermosas y abundantes flores, ademas del equilibrio
que proporcionan al paisaje.

Por ser las plantas ornamentales poco estudiadas en el estado de Nuevo
Le6n y en particular para el drea metropolitana de Monterrey, se consideréd
pertinente realizar un inventario de éstas plantas con el fin de ampliar el
conocimiento de las mismas y aportar datos que contribuyan a estudios
posteriores.

Algunos autores han trabajado diversos aspectos acerca de la Flora en el

SIDA 18(2): 579-586. 1998

580 Sipa 18(2)

estado de Nuevo Leon, sin embargo ninguno de ellos, tiene un enfoque
particular hacia las plantas ornamentales, aunque algunos por su amplitud
hacen mencion de algunas de ellas. Entre estos trabajos destacan el de Gonzalez

1888), pionero oa 108 estudios botanicos en el estado, él realizé un inventario
floristico acerca d t 1 Nuevo Leén, encontrando
cerca de 264 especies incluidas en 70 familias. Por su parte Rojas-Mendoza
(1965), describié la vegetacién y la flora para el Estado, él reporta 148 familias,
657 géneros y un total de 1484 plantas con categoria de especie, subespecie,

variedad o forma.

Es conveniente mencionar que si existen trabajos florfsticos o inventarios
de plantas ornamentales para otras ciudades en el pafs, como es el caso de
Teresa de Tovar (1982), quién llevé a cabo un estudio descriptivo de los
arboles y arbustos mas comunes en el Bosque de Chapultepec, describiendo
un total de 57 familias, 113 géneros y 168 especies. Capitanachi-Moreno y
Amante-Haddad (1995), realizaron un estudio de vegetacién urbana en Xalapa,
Veracruz, encontrando 443 especies correspondientes a 317 géneros incluidos
en 117 familias.

AREA DE ESTUDIO

E] area metropolitana de Monterrey comprende los municipios de San
Nicolas de los Garza, Guadalupe, San Pedro Garza Garcia, Santa Catarina
y Monterrey. merase una eer de 1976.480 Km’, localizandose entre
las coordenadas 25° 25° 47' de latitud Norte y entre los 100° 06' y
100° 43’ de longitud on con una altitud de 554 msnm. La precipitacién
pluvial es escasa, oscilando la media general anual entre 300 y 600 mm,
pero hay zonas que registran Iluvias anuales mayores de 800 mm.

MATERIAL Y METODO

La colecta de las plantas se llevé a cabo en plazas y jardines publicos del
area metropolitana de Monterrey, las cuales suman un total de 393, debido
a su considerable numero, se hizo necesario calcular un tamafio de muestra
estadisticamente significativo, para lo cual se realiz6 un muestreo piloto,
en tres plazas o jardines publicos (seleccionados aleatoriamente) de cada
municipio y con los resultados obtenidos se realizo el calculo para el tamafio
de muestra total (numero de plazas y jardines a muestrear) mediante la formula
de muestreo estratificado (Abad de Servin y Servin-Andrade 1978), lo cual
nos permitid conocer el numero de plazas y jardines a muestrear en cada
municipio, asf como el numero total para el drea metropolitana de Monterrey.

El tamafo de muestra total fue de 52 plazas y jardines, de las cuales cinco
corresponden al municipio de Santa Catarina, tres al municipio de Guadalupe,
ocho al municipio de San Pedro, dieciséis al municipio de San Nicolas y
veinte al municipio de Monterrey.

Rocua, et al., Flora ornamental de Monterrey 581

El material bioldgico obtenido durante las colectas fue debidamente
herborizado y etiquetado para su posterior identificacién, siguiendo el criterio
de Bailey (1949), Correll y Johnston (1970) y Elias (1980), considerando
ademas los trabajos de Martinez (1979), Sanchez-Monje (1980), Capitanachi-
Moreno y Amante-Hadadd (1995). Los ejemplares se encuentran depositados
en el Herbario de la Facultad de Ciencias Bioldégicas, U.A.N.L. (UNL).

RESULTADOS

Se colectaron un total de 137 especies, las cuales estdn incluidas en 68
familias representativas de las plantas ornamentales. La familia con el nimero
mayor de especies corresponde a Leguminosae con 10 especies, seguida por
las familias Liliaceae y Malvaceae con 8 y 6, las familias Apocynaceae, Compositae
y Euphorbiaceae con 5 especies y el resto de las familias con menor nimero
de miembros (Fig. 1 y listado).

Las especies que con mayor frecuencia se encontraron en los sitios de
colecta fueron: Fraxinus sp. (80.76%), Ligustrum lucidum Ait. (59.61%),
Rosa centifolia L. (42.30%) y Melia azedarach L. (36.52%)

Big. 2

Se aprecié que las plantas encontradas en las plazas y jardines publicos
del drea metropolitana de Monterrey son basicamente hierbas (37.95%),
arboles (30.65%), arbustos (26.27%), palmas (2.91%) y trepadoras (2.18%)
(Cuadro 1).

En relacién al origen de la flora ornamental del area en estudio, se encontré
que del total de especies aproximadamente un 38% son especies nativas y
el resto (62%) son especies introducidas provenientes bdsicamente de Europa,
Asia y Africa.

Familias

No. de especies

Fic. 1. Familias con mayor numero de especies amg en los parques y jardines
Ma P
publicos del 4rea metropolitana de Monterrey, N.L., Méxic

582 SIDA 18(2)

Por otra parte en el Cuadro 2 puede apreciarse que existen diferencias
entre la diversidad florfstica presente en los municipios del area metropolitana
de Monterrey, donde sobresale Monterrey con un promedio de 19 especies
por area verde, en tanto que el municipio de Guadalupe solo existen 6 especies
en promedio.

DISCUSIONES Y CONCLUSIONES

De las especies colectadas por Gonzalez (1888) para el Estado de Nuevo
Leén, 25 coinciden con las aqui encontradas; asi como 73 con el inventario
realizado por Rojas-Mendoza (1965) para el estado y 24 con Teresa de Tovar
(1982) quien estudid el bosque recreativo de Chapultepec; con el estudio
realizado por Capitanachi-Moreno y Amante-Haddad (1995) coincide con
45 especies reportadas en este trabajo

La forma bioldégica mas frecuente fue la herbacea, esto contrasta con el
estudio realizado por Tovar de Teresa (1982) en el Bosque de Chapultepec,
ya que ella menciona que las especies arboreas son las que tienen mayor
diversidad y frecuencia. Por su parte Capitanachi-Moreno y Amante-Haddad
(1995) concuerda con la forma herbacea.

En relacién al origen de las especies se encontré que solo un 38% son
especies nativas y el resto son introducidas (62%), lo cual es congruente
con lo mencionado por Rzedowski y Calderon de Rzedowski (1990), quienes
mencionan que dentro del conjunto de plantas introducidas en México la
mayoria tiene un origen europeo, siendo f la procedencia
mediterranea.

Frecuencia (%)

F. sp. L krckatem

8 gana W pment
Especies

Fic. 2. Especies ornamentales mds Vie en los parques y jardines publicos del area
metropolitana de Monterrey, N.L.,

Rocna, et al., Flora ornamental de Monterrey

ae 1. Formas bioldgicas de las pl

583

tales del drea metropolitana de Monterrey, N.L.,

Méxicc

Forma bioldgica

No. especies

Porcentaje

Hieba 52 37.95

rbol 42 30.65
Arbusto 36 26.67
Palma 4 2.91
Trepadora 3 2.16
Cuadro aR W134 4 2 1% yea 1] ] dahd

N.L., México.

Municipio No. especies promedio No. minimo No. maximo
Monterrey 19 1 70
San Nicolas 14 1 43
Guadalupe 6 1 13
San Pedro 115 20
Santa Catarina 15 7 29

APPENDICE
Listado de especies ornamentales en parques, plazas Y jardines publicos del
drea Metropolitana de Monterrey, N.L., México.

ACANTHACEAE

Ruellia brittoniana Leonard, NH
ADIANTA

Pteris ae L., IH
AIZOACEAE

Sesuvium portulacastrum L., TH
AGAVACEAE

Agave asperrima Jacobi, NH

Sansevieria zeylanica Willd., 1H

Yucca carnerosana (Trel.) McKelvey, NP
AMARANTHACEAE

Alternanthera repens (L.) Kuntze, TH

Amaranthus caudatus L., TH
AMARYLLIDACEAE

Crinum zeylanicum L., [IH

Hippeastrum equestre Herb., TH

Nothoscordum bivalve (L.) Britton, IH
ANACARDIACEAE

Rhus laurina Nutt., IA

APOCYNACEAE
Carissa grandiflora A. DC., [Ar
Nerium oleander L., VAr
Ochrosia elliptica Labill., I
Thevetia peruviana (Pers.) K. Schum., IA
Vinca rosea

ARACEAE

Monstera delictosa Liebm., NH

Philodendron cordatum Kunth, IH
ARALIACEAE

Hedera helix L., YT
BEGONIACEAE

Begonia semperflorens Link & Otto, IH
BIGNONIACEAE

Chilopsis linearis Sweet, NA

Jacaranda mimosifolia D. Don, IA

Tecoma stans (L.) Juss ex HBK, NAr
BORAGINACEAE

Cordia boissieri A. DC.,

NAr
Ebretia anacua (Teran & Berl.) I.M. Johnst,
NA

584
CANNACEAE

Canna indica \.., WH
CARICACEAE

Carica papaya L., NA
CAPRIFOLIACEAE

Abelta chinensis R. Br., [Ar
Lonicera japonica Vhunb., IT
Sambucus mexicana Presl, NA
CARYOPHYLLACEAE
Dianthus caryophyllus L. ex J. R. & G. Forst.,
IH

CASUARINAC
Casuarina equisetifolia L., LA
COMMELINACEAE
Commelina tuberosa L., NH
Tradescantia virginiana L., NH
COMPOSITAE
Artemisia albula Nutt., NH
Cosmos bipinnatus Cav., NH
lagetes erecta L., NH
Tagetes patula L., NH
Zinnia elegans Jacq., NH
CONVOLVULACEAE
Ipomoea vo“ Mart., NT

Kalanchoe eee Poellnitz, [TH
CUPRESSACEAE
Cupressus sempervirens L., IA
Juniperus monosperma (Engelm.) Sarg., NA
die occidentalis L., N
CYPE
S ra eile Ly LEE

ERICACEAE
Arctostaphylos glauca Lindl., NAr
Rhododendron mucronatum Don, LAr

EUPHORBIACEAE
Cnidoscolus chayamansa Mc. Vaugh., NAr
Croton ciliato-glandulosum Ort., NAr
Euphorbia pulcherrima Willd. ex Klotzsch,
NAr
Jatropha dioica Cerv., NAr
Sapium sebiferum (L.) Roxb., IA
FAGACEAE
Quercus virginiana Mill., NA

SIDA 18(2)

FOUQUIERIACEAE
Fouquieria splendens Engelm., NAr
GERANIACEAE
Pelargonium zonale Ait., 1H
GRAMINEAE
Bambusa vulgaris Schrad. ex Wendl., -H
Cortaderia selloana Aschers & Graebn., [H
Saccharum officinarum |
Stenotaphrum secundatum (Walt.) O.
Kuntze, TH
HAMAMELIDACEAE
Liquidambar styraciflua L., NA
JUGLANDACEAE
Carya cordiformis (Wangenh.) K. Koch,
NA

LABIATAE
Ocimum basilicum L., TH
Teucrium cubense L.,

LAURACEAE
Persea americana Mill., NA

LEGUMINOSAE
Acacia farnesiana (L.) Willd., NA
Bauhinia variegata L
Caesalpinia mexicana Gray., NAr
Caesalpinia pulcherrima (L.) Swartz, -Ar
Erythrina coralloides D.C., NA
Leucaena glauca L., NAr
Mimosa matacophylla Gray, NH
Parkinsonia aculeata L., LAr
Prosopis glandulosa Vorr., NA
Trifolium repens L., UH

LILIACEAE
Aloe vera L., TH
Allium scaposum Benth., NH
Asparagus sprengert Regel., IH
Asparagus virgatus Baker, 1H
Chlorophytum capense Vhunb., 1H
Dasylirion longissimum Lem., UH
Hemerocallis middendorffii Vrauty., 1A
Ophiopogon jaburan Lodd., TH

LYTHRACEAE
Lagerstroemia indica L., IA

MAGNOLIACEAE
Magnolia grandiflora L., IA

MALVACEAE

Hibiscus rosa-sinensis L., Ar

Rocna, et al.,
Hibiscus syrtacus L., TAr
Lavatera trimestris L., [Ar
Matlvaviscus arboreus Cav., NAr
Sida abutifolia Miller,

Thespesia populnea (L.) Soland ex Correa,
[Ar

MELIACEAE

Melia azedarach L., 1A
MORACEAE

Morus alba L.

Ficus elastica Roxbg., IA
MUSACEAE

Musa paradisiaca L., YAr
MYRTACEAE

Callistemon specious DC., NAr

Eucalyptus globulus Labill., IA
NYCTAGINACEAE

Bougainvillea glabra Choisy, LAr

Mirabilis jalapa L., NH
OLEACEAE

Fraxinus velutina Torr., NA

Ligustrum japonicum Thunb., [Ar

Ligustrum lucidum Ait., LA
PALMAE

Arecastrum romanzoffiana Becc., IP

Phoenix dactylifera L.,

Washingtonia filifera (Linden) H. Wendl.,
NP

PINACEAE

Pinus halepensis Mill., NA
PITTOSPORACEAE

Pittosporum tobira Ait., LAr
PLATANACEAE

Platanus occidentalis L., NA
PLUMBAGINACEAE

Plumbago capensis Thunb., [Ar
POLYPODIACEAE

Nephrolepis exaltata (L.) Scott, NH
PUNICACEAE

Punica granatum L., lAr
RANUNCULACEAE
Delphinium ajacis L., UA

RHAMNACEAE
Colubrina greggii Wats., NAr

Flora ornamental de Monterrey

585

ROSACEAE
Eriobotrya japonica Lindl., IA
Prunus persica (L.) Osbeck., IA
Pyracantha coccinea L., lAr
Rosa centifolia L., LAr
RUBIACEAE
Gardenia jasminoides Ellis, [Ar
RUTACEAE
Citrus aurantifolia (Christm.) Swingle,
IA

—

Citrus sinensis Osbeck., IA
Sargentia gregeii Wats., NAr
Zanthoxylum fagara (L.) Sarg., NAr
SALICACEAE
Populus tremuloides Michx., 1A
Salix nigra Marsh., 1A
SAPINDACEAE
‘oelreuteria paniculata Laxmn., IA
SCROPHULARIACEAE
Leucophyllum frutescens (Berl.) 1.M.
Johnst., [Ar
Mimutlus puniceus Steud., [Ar
SOLANACEAE
Petunia axillaris BSP., TH
Petunia hybrida Hort. ex Vilm, TH
STRELITZIACEAE
Strelitzia reginae Banks, TH
TAMARICACEAE
Tamarix gallica L., 1A
TAXODIACEAE
Taxodium mucronatum Ten., NA
TROPAEOLACEAE
Tropaeolum majus L., WH
ULMACEAE
Celtis laevigata Willd., NA
Celtis pallida Torr., NA
UMBELLIFEREAE
Coriandrum sativum L., TWH
bs ean ts
Lantana camara
Verbena Hiern Nutt. IH

: las siglas que aparecen a la derecha
corresponden al origen: - desconocido, N nativa,
] introducida y ala a bioldgica: H herbdcea,

A arbol, Ar arbusto, P palma, T trepadora.

586 Stipa 18(2)
REFERENCIAS

Apap De Servin, A. y L. A. SERVIN-ANDRADE. 1978. Introduccién al muestreo. Primera
edicién. Editorial Limusa, México, D.F. Pp. 105-141.

Baitey, L.H. 1949. Manual of aleeied plants. MacMillan Publishing CO, Inc, New York.

CAPITANACHI-MoRrENO, C.Y.S. AMANTE-Habpab. 1995. Las areas verdes urbanas en Xalapa,
Veracruz. Catalogo de Flora Urbano. Universidad Veracruzana/Instituto de Ecologia A.C,

CorretL, D.S. and JOHNSTON, M. 0. Manual of the vascular plants of Texas. Pub-
lis ed Texas Research ie aaacon: Renner Texas.

Eutas, S.T. 1980. Trees of North America. ee Nostrand. Reinhold Company, New York.

ee J.E. 1888. Un discurso y un cat Clasificadas. Imprenta Catdlica.

Martinez, M. 1979. Catalogo de ee ete sie de plantas Mexicanas.
Primera edici6n. Fondo de Cultura Econédmica. Méx

RzEDOSWKI, J. y G. CALDERON DE RZEDOWSKI. 1990. oe see el elemento africano en la

Jora adventicia de México. Acta Bot. Mexicana. 12:21—

BOS ME poe P. 1965. Generalidades sobre la ea del estado de Nuevo Leon y

u flora. Tésis doctoral inédita Universidad Nacional Auténoma de México,

a

o, DLE
Sancti Mow E. 1980. Diccionario de plantas agrfcolas. Ministerio de Agricultura. Madrid,
Espa
‘TERESA Dr Tovar, L. 1982. Estudio descriptivo de los arboles y arbustos mas comunes del
1e de C are ee ‘Tesis profesional. Facultad de Ciencias. Universidad Nacional
a ae de Méx

DOCUMENTED CHROMOSOME NUMBERS
1998:1. CHROMOSOME NUMBERS IN CAREX
SECTION OVALES (CYPERACEAE): ADDITIONS,
VARIATIONS, AND CORRECTIONS

PE. ROTHROCK

Randall alias Center
Laylor University
Upland, IN eas 1001, U.S.A.

Ako REZNICER

University of Michigan Herbarium
North University Building
Ann Arbor, MI 48109, U.S.A.

ABSTRACT

Ch ts for 14 species of Carex section Ovales are d ted completing
at least a single report for each of the over 40 species known for eastern North America.
The haploid numbers in this report ranged from 7 = 24 to n = 42. Counts for Carex feta,
C. muskingumensis, C. oronensis, and C. tetrastachya det ined for the first time and
six aneuploid (sensu lato) series are indicated. The only previously published counts for C.
adusta, C. argyrantha, and C. foenea (C. aenea) were found to be incorrect.

RESUMEN
Se documentan los recuentos de 14 especies de Carex seccié6n Ovales completando al
menos una cita de las mas - 40 especies nas del este de América del Norte. Los
numeros haploides varfan de 24 hasta » = 42. Se hacen por primera vez recuentos de
Carex feta, C. muskingumensts, Cx OYONENSIS, Y C tetrastachya y se indican seis series aneuploides
ages Los tinicos recuentos previos de C. adusta, C. argyrantha, y C. foenea (C. aenea)
e ha encontrado que son incorrectos

INTRODUCTION

Carex section Ovales is a large, coherent group of sedges with a center of
diversity in North America. At least 40 species occur east of the Rocky Mountains
and about as many species occur from the Rocky Mountains westward
(Mastrogiuseppe et al. in review). Numerous karyological studies over the
past 50 years (Whitkus 1991) have shown much chromosomal variation at
the sectional, species, and population level. Even individual plants may have
some cell to cell changes in karyotype. Much of this variation is thought to
be the consequence of polycentric chromosomes with diffuse centromeres
(Grant 1981). Polycentric chromosomes have the ability to fragment or fuse
with other chromosome pieces and still retain the potential of completing

SIDA 18(2): 587-592. 1998

588 Stipa 18(2)

normal movements during mitosis and meiosis. In spite or because of this
variation, chromosome data have been informative in understanding the
taxonomy and probable phylogenetic relationships among these species
(Rothrock & Reznicek 1996a). And previously unrecognized ier such
as Carex ozarkana and C. molestiformis, have been revealed to have distinc-
tive chromosome numbers (Rothrock & Reznicek 1996b; Reznicek & Rothrock
199

As part of ongoing taxonomic studies of eastern North American spe-
cies of Carex section Ovales and especially as part of the Flora of North
America project, an effort was made to complete a karyological survey of
all Species: ie occur in | eastern sone America and, where possible, to
variability. This update documents the karyotypes

of 40 plants see 14 species.

MATERIALS AND METHODS

As described previously (Rothrock & Reznicek 1996a), living plants were
collected in the field during their fruiting stage and cultivated in a sand-
peat- perlite medium under greenhouse conditions. In this study, two spe-
cies, C, praticola and C. xerantica, were grown from seed taken from her-
barium specimens. Cold treatment was used to stimulate the production
of new inflorescences by the following spring. pe spikes were pre-
served in methanol, chloroform, propionic acid (6:3:2). Within the subse-
quent 72 hours, anthers were dissected from the file and squashed in
2% lactic-acetic-orcein (Cooperrider & Morrison 1967). Meiosis I chro-
mosome figures were examined from five or more pollen mother cells.
Photographs and drawings were made with a Nikon Labophot-2 micro-
scope using phase contrast at 1000x magnification.

Voucher specimens were typically prepared at the time of field collec-
tion or occasionally from fruiting greenhouse material. Vouchers, on de-
posit at the University of Michigan Herbarium (MICH), include photo-

micrographs and drawings of countable figures.

RESULTS AND DISCUSSION

The chromosome counts for the 14 species reported in this study ranged
from 7 = 24, the lowest now known for section Ovales (Whitkus 1991), to

= 42 (Table 1). For six of the species intraspecific variation in chromo-
some number is reported. Five counts published by Love and Love (1981)
are clearly at odds with our results and, given the often subtle differences
between species, likely indicate faulty identification of the plant being ex-
amined.
Carex adusta EF. Boott.—Our material from Washington Co., Maine, comes
from near the eastern extreme of the distribution of C. adusta. Its n = 39

ROTHROCK AMD REZNICEK, Chromosome numbers in Carex

589

Tape 1. New and published chromosome numbers for select species of Carex section Ovales.

Species

Location: Vo

ucher or Reference

Carex adusta F. Boott

Carex argyrantha Tuckerman

Carex brevior (Dewey) Mackenzie

Carex feta L.H. Bailey
Carex foenea Willd.

Carex hyalina ¥. Boott

Carex longii Mackenzie

Carex merritt-fernaldii Mackenzie

Carex molesta Bright

Carex muskin gUMeNSIS Schwein.

d

Carex oronensis Ferna

Carex praticola Rydb.

Carex tetrastachya Scheele

Carex xerantica L.H. B ailey

A hapa es - 2 oe A.A.R 10560
ive & ee
Hancock Co., ME: PE.R. 3479
Washin ton Co., aa A.A.R. 10087
entre oe ,PA 3494
Wahl 1940

r éve & Love 1981

Kaufman Co.,

pees

Lane Co.,

. ancock Co.,

Tx: A.ALR. 10345b
ALR.

OR: Wilson 7869

ME: Ce S ee
JINEE

ten cock Co., Oa on Sr

rafford Co.
Love & Love
Morris Co.,

Rothrock &

Cumberlar
Rothrock &
Rot ee &

Suetor

TX: A 8489
each 1996a

id Co., a ne R. 3638

Rez 996a
ae i
a PE.R. 3475
981

vee i 1 woe

Bolivar Co.,

MS: Bryson 12209

White Co., IN: RE.R. 3356

Grant Co., IN
Macon Co.
Wahl 1940

Huntington

Hancock Co., M
Penobscot Co.,

Thunder Ba
Packer &
Whit

: PELR. 2482
, MO: PE.R. 3567.5

1940
Love & Léve 1981

Co., IN: PE.R. 2132

E: PE.R. 3481
ME: PE.R. 3483

ee waaae Oldham 17731a
a tku

ve & ee ee

Kaufman Co.,
Zav

jee a

TX AAR. 10344

maw AAR.

andes poe aaice Ontario: Oldham 17732

e & Lov

* problematic, unvouchered reports; see text

590 Sipa 18(2)

disagrees with the 7 = 32 listed by Léve and Léve (1981). Unfortunately
many Léve and Léve chromosome counts for Carex section Ovales, including
this one, have no known, extant voucher specimens. We suspect, however,
that their 7 = 32 count may be based upon an individual belonging to some
other species, perhaps C. brevior (Dewey) Mackenzie.

Carex argyrantha Tuckerman.—Based upon single plants from 3 locali-
ties, this species had a consistent 7 = 40. Wahl (1940) published a count of
n= 40 for this species under the misapplied name C. foenea. Love and Love
(1981) incorrectly attribute an 7 = 32 to this species. Their material came
from a Manitoba source which lies outside the range of C. argyrantha.
Unfortunately no voucher is known for their count.

Carex brevior (Dewey) Mackenzie.—Our material (10 plants) gave a sur-
prisingly broad range of chromosome counts, from 7 = 24 to 32. Morpho-
logically the species is variable but we were unable to correlate any particu-
lar phenotype with chromosome count. We thus have no reason as yet to
suspect that 2 or more sibling species are hidden within this taxon. The
lowest counts, 7 = 24, 24 + IV, and 26 are from unusually robust individu-
als collected in Kaufman Co., Texas. The 2 = 30 individual was notewor-
thy in having clear nervation on the dorsal face of the perigynia. A previ-
ous report for Carex brevior (with voucher specimen documentation available)
isn = 34 (Love & Love 1981). This report is plausible, though the number

remains unconfirmed.

Carex feta L.H. Bailey.—This species, from western North America, had
an # = 33. Given the size and shape of the perigynia and achenes as well as
the clear collar at the orifice of the leaf sheath, we suspect that C. feta is
most closely related to C. festucacea Schk. ex Willd. found in eastern North
America. The later species has a chromosome range of 7 = 33 + III, 34, and

35 (Wahl 1940; Rothrock & Reznicek 1996a).
Carex foenea Willd.—Our material (4 plants) had ” = 41 and 42. In fact,

the short aneuploid series was observed within the population from Hancock
Co., Maine. Many taxonomic treatments (e.g. Fernald 1950) have incor-
rectly called this taxon C. aenea Fern. Love and Léve (1981) published an
n = 32 tor C. aenea for material collected from near Brandon, Manitoba.
No voucher specimen from this locality could be found. On the other hand,
their voucher from Maskwa rapids, Manitoba (Love & Léve 5512, WIN!)
is a specimen of C. foenea and the annotation label indicates that the chro-
mosome count was “27 = 64.” Unfortunately this voucher was intended
for their report of C. tenera, a species with 2 = 26 to 28 (Wahl 1940; Rothrock
& Reznicek, unpublished data).

The relatively high chromosome number for C. foenea compares favor-
ably to that of several other species possessing long pistillate scales. These

ROTHROCK AMD REZNICEK, Chromosome numbers in Carex 591

include C. adusta (n = 39), C. argyrantha (n = 40), and perhaps C. silicea
Olney (7 = 37, 38).

Carex hyalina F. Boott.—This specimen from the southwestern extreme
of the species’ range had an 7 = 37. Our previous report (Rothrock & Reznicek
1996a) for the rare C. Ayalina, from the northern portion of its range in
Mississippi, also provided an x = 37.

Carex longii Mackenzie.—Previous work (Rothrock & Reznicek 1996a)
found 7 = 28 + IV and 31 for plants from the coastal plain of the south-
eastern US and from southern Michigan. This plant with n = 29 comes
from the Cumberland Mountain region of Tennessee.

Carex merritt-fernaldii Mackenzie.—Given a count by Tanaka (1942) of
n = 35, our result of 7 = 37 suggests possible aneuploidy for this species.
Love and Léve (1981) offer an undocumented count of 27 = 68. However
their count must be in error since the plant material came from Melita in
extreme southwestern Manitoba, well outside the known range for C. merritt-
fernaldii in that province.

Carex merritt-fernaldii often is morphologically confused with C. brevior.
The differences in chromosome condition, however, reinforce its recogni-
tion at the species level.

Carex molesta Bright—Previous work found » = 34 for this taxon (Wahl
1940). Love and Léve (1981) also found 2 = 34 for undocumented mate-
rial from Manitoba where, as far as we know, C. molesta does not occur. We
confirm 7 = 34 but also found an aneuploid variation of 7 = 35.

Carex muskingumensis Schwein.—The count of = 40 (based upon 2
plants) is the first for this species. For a member of Carex section Ovales, C.
muskingumensis is quite distinctive. The stiff inflorescence, spikes, and lan-
ceolate perigynia are unusually large. The vegetative culms have numerous
evenly spaced leaves reminiscent of the C. tribuloides group (with n = 32 &
35), the mostly likely near relative of C. muskingumensis.

Carex oronensis Fernald.—The counts of 7 = 37 are the first for this rare
endemic of the Penobscot River valley in central Maine. Dibble (1991)
hypothesizes that the European C. ovales is a near relative. However, this is
not supported by the numerous chromosome counts for C. ovalis which
range from 7 = 32 to 34 (Whitkus 1991). We are unable to suggest pos-
sible close affinities between C. oronensis and other species in eastern North
America.

Carex praticola Rydb.—Primarily a species of western North America, this
plant comes from a disjunct southern outlier of the species’ natural distri-
bution. Its 7 = 39 is identical to that of Packer and Whitkus (1982) and
Whitkus (1991), who additionally reported 7 = 38. The unvouchered re-

592 Sipa 18(2)
port of 7 = 32 by Léve and Love (1981) is highly suspect.

Carex tetrastachya Scheele.—Formerly known as C. brittoniana, C. tetrastachya
displays an aneuploid series. Among the material available, we observed x
= 30+ HI (1 plant), 32 (3 plants), 33 (4 plants). Correll and Johnson (1970)
suggest that this taxon and C. /yalina are most closely related. Vegetatively
we ey find the two to be quite distinctive and the C. hyalina counts
of 2 = 37 do not immediately suggest a close affinity.

Carex xerantica L.H. Bailey — Our plant from a disjunct and eastern out-
lier of the species in Ontario yielded an 7 = 34. This confirms an unvouchered
(but geographically plausible) report by Love and Léve (1981) for this northern
species.
ACKNOWLEDGMENTS

Our thanks are given to Charles Bryson, Garrett Crow, Alison Dibble,
Stanley Jones, and J.K. Wipff for providing living material or assistance in
the field; to Bruce Ford and William Weber for invaluable herbarium in-
formation; and to Michele Jordan for assistance in the greenhouse.

REFERENCES

CooperribDER, [S. and J.H. Morrison. 1967. Lactic-acetic-orcein as a chromosome stain.
Michigan Bot. 6:176-178.

Corre, D.S. and M.C. JOHNSON. 1970. Manual of the vascular plants of Texas. Texas
Research a ation, Renne

Dipsie, A.C. . Carex oronensis : Lyperaceae), a possible neoendemic of Maine. M.S.
thesis, eee of Maine, Oron

FERNALD, M.L. 1950. Gray’s manual of botany. Reprinted edition. D. Van Nostrand Co.,
New York, NY.

GRANT, V. 1981. Plant speciation (second edition). Columbia University Press, New York,
N

Love, A. and D. Love. 1981. Chromosome number reports LXXIII. Taxon 30:829-861.

Maestrociusepre, J. RE. Rorurock, P. BALL, A.C. DipBLe, and A.A. REZNICEK. Carex sec-
tion Ovales (Knuth) Christ, Flora of North America (In review).

Packer, J.G. and R. Wirrkus. 1982. Chromosome number reports LXXV. Taxon 31:342—
368.

REZNICEK, A.A. and PE. RoTHROcCK. 1997. Carex molestiformis (Cyperaceae), a new spe-
cles . section enn from the Ozark Mountain region. Contr. Univ. Michigan Herb.
21:299-308

RorHrock, P.E. and A.A. REZNICEK. 1996a. Chromosome numbers in Carex section Ovales
(Cyperaceae) from eastern North America. Sida 17:251-258.

Rorurock, PE. and A.A. Reznicek. 1996b. A new species of Carex section Ovales (Cyperaceae)
occurring in the Ozark Mountain region. Brittonia 48:104-1

TANAKA, H. 1942. Chromosome studies - ee XX. Chromosome numbers of Carex

eee ee a and Biol, 2:220—2

0. Chromosome a and meiosis in the genus Carex. Amer. J. Bot.

Rie

07: 458- 47(
Wuitkus, R. 1991 Chromosome counts of Carex section Ovales. Bot. Gaz. (Crawfordsville)
5 30.

VASCULAR PLANTS NEW TO KANSAS

CRAIG C. FREEMAN, c-freeman@ukans.edu
RONALD L. MCGREGOR
CALEB ru MORSE, cmorse@falcon cc.ukans.edu
RL. McGregor Herbarium, Division of Botany
Natural History Museum and Biodiversity Research Center
University of Kansas, 2045 Constant Avenue
Lawrence, KS 66047-3729, U.S.A.

Recent field and herbarium studies have yielded 37 vascular plant taxa pre-
viously undocumented in Kansas. Seventeen of these are introduced and
naturalized; the others are considered native to Kansas. Notes about these
taxa are provided to update distribution data contained in the Afdas of the
Flora of the Great Plains (1977) and the Flora of the Great Plains (1991) and
to make these data more widely available to botanists working on the Flora
of North America project. All voucher specimens are deposited in the R.L.
McGregor Herbarium (KANU), University of Kansas, unless otherwise
indicated.

Acalypha deamii (Weath.) H.E. Ahles (Euphorbiaceae).—Acalypha deamii
occurs at scattered localities from the Ohio River valley west to Kansas. It
typically grows in floodplain forests along rivers and streams. Geoffrey A
Levin, Ilinois Natural History Survey, discovered a single sheet among KANU
specimens of Acalypha tating material for the Flora of North America
project in 1995. The collection site, in Chase County, was visited in 1995
but no A. deamii was found. A population of several dozen plants was dis-
covered the following year in east-central Shawnee Co.

Voucher specimens: KANSAS. Chase Co.: Cottonwood Falls, 6.5 mi S, nae area
along Rock Creek, moist, rich soil, 12 Sep 1975, Stephens 88044. Shawnee Co. 25
RIGE, sec 1, W 1/2, Tecumseh, 0.5 mi S, 0.5 mi W, elev 880-920 ft, flood Shih ote
with scattered limestone outcrops along Stinson Creek between US 40 & | 50, 27 Aug

1996, Freeman 8172.

Alyssum desertorum Stapf var. desertorum (Brassicaceae).—This small,
introduced annual is an innocuous range weed throughout much of the
western U.S. (Rollins 1993). In the northern Great Plains it is encountered
on disturbed prairies, along roadsides, and in waste places, especially in the
High Plains (Great Plains Flora Association 1991). A single population was
discovered in northwest Kansas, where plants occupy a cemetery and shoulder
of an adjacent gravel road.

SIDA 18(2): 593-604. 1998

594 SipA 18(2)
Voucher specimen: KANSAS. Thomas Co.: 17S, secre sec 28, NW 1/4, NW 1/4,

Gem, 0.25 mi SW, Gem Cemetery, elev 3090 ft, mowed, upland cemetery with some remnant

shortgrass prairie, 24 Apr 1996, Freeman 7939

Amaranthus californicus (Mog.) $. Watson (Amaranthaceae).—A num-
ber of sheets of this species recently were discovered among KANU speci-
mens identified as A. blitoides S. Watson. Amaranthus californicus can be
distinguished from A. b/itoides by its smaller seeds (0.8—1.1 mm diameter
vs 1.2—1.7 mm diameter) and fewer (1—3 vs. 4—5), ee. pistillate se-
pals (Henrickson 1993). California pigweed 1s a widespread native in the
western U.S. and first was reported in the Great Plains based on a 1995
collection from Dawes Co., Nebraska, by Steve Rolfsmeier. We also have
vouchers from South Dakota (Shannon Co.) and Texas (Deaf Smith, Gray,
and Parmer cos.)

Voucher specimens: KANSAS. Barton Co.: Great Bend, 7 mi NE, area of Cheyenne
Bottoms, oo dry pond shore, 15 Aug 1977, McGregor 30538. Morton Co.: Elkhart, 6

ni N, 3 mi W, edge of dried pool, 13 Aug 1982, McGregor 33603. Rush Co.: Bison, | mi
S. bank of Sand Creek, 15 a 1977, McGregor 30541. Sherman Co.: Goodland, 9 m1 S,
3 mi W, Sherman County Lake, along lake shore, 15 Aug 1983, McGregor 34742.
Antennaria parvifolia Nutt. (Asteraceae). —Small-leaf pussy's-toes is a common,
widespread, spring ephemeral in the northern and central Great Plains (Great
Plains Flora Association 1991). Earlier collections document its range in
the central Great Plains south to the near the north border of Kansas. The
species is added to the Kansas flora based on two collections from the northwest
part of the state.

Voucher specimens: KANSAS. Norton Co.: 15S, R24W, sec 23, SE 1/4, SE 1/4, NW
1/4, Lenora, SE edge of town in Lenora Cemetery, elev 2310-2320 ft, hill above Lost Creek,
overlooking the floodplain of the North Fork Solomon River, site with sce mixed
grass prairie vegetation, 8 May 1997, Freeman 8786; 15S, R22W, sec 8, W 1/2, W 1/4,
NE 1/4, Edmond, 1 mi N, Edmond Cemetery, eley 2220 ft, level, upland, san cem-
etery E of draw dominated by mixed grass prairie, 8 May 1997, Freeman 8795.

Arabis hirsuta (L.) Scop. var. adpressipilis (M. Hopkins) Rollins (Brassicaceae).—
Two varieties of hairy rockcress occur in the Great Plains. Arabis hirsuta
var. pycnocarpa (M. Hopkins) Rollins has trichomes on the stem spreading
and unbranched and occurs throughout the northern Great Plains south-

ward into central Kansas. Variety adpressipilis, with appressed and forked
stem trichomes, barely enters the southeastern Great Plains. It is the com-
mon variety throughout Missouri (Steyermark 1977) and occurs in extreme
southeast Kansas at a single locality.

Voucher specimen: KANSAS. Cherokee Co.: T34S, R25E, sec 35, NW 1/4, NE 1/4 &
NE 1/4, NW 1/4, jct US 166 & KS 26, 1.5 mi N, Schermerhorn Park, elev 830-900 ft,
rocky, oak- ee ee on bluffs and uplands N of Shoal Creek, scattered along creek in
NW part of park, 23 Apr 1992, Freeman 4323.

Armoracia lacustris (A. Gray) Al-Shehbaz & V.M. Bates (Brassicaceae).—
Al-Shehbaz and Bates (1987) mapped the distribution of this aquatic pe-

FREEMAN et al., Plants new to Kansas 595

rennial throughout the Great Lakes region, Midwest, lower Mississippi River
valley, and at other widely scattered sites in the eastern U.S. A population
discovered in southeast Kansas extends the western range of the species roughly
30 kilometers. Unfortunately, it was destroyed by construction activity short
after it was discovered.

Voucher specimen: KANSAS. Neosho Co.: just S of South Mound in shallow body of
water by road near railroad, 25 Jun 1995, Holland 8358.

<=

Aster lateriflorus (L.) Britton var. lateriflorus (Asteraceae).—This aster
erroneously has been attributed to Kansas (Gates 1940; Great Plains Flora
Association 1991; Gleason & Cronquist 1991) based on misidentified specimens
of A. ontarionis Wiegand. It is fairly common in mesic forest and wood-
lands in Missouri (Steyermark 1977), anda population discovered during
recent field studies in northeast Kansas confirms the species occurs in Kansas.

Voucher specimen: KANSAS. Leavenworth Co.: T8S, R22E, sec 15, SE 1/4, Ft. Leavenworth
Military Reservation, W-central part, slopes ESE of Wagner Point, elev 900-1050 ft, mesic,
oak-hickory-maple-basswood forest and small, xeric, tallgrass prairie remnants on ridge E

of Sheridan Drive, 3 Oct 1995, Freeman 7808

Atriplex prostrata Bouchér ex DC. (Chenopodiaceae).—An examination
of KANU Artriplex specimens by Steve Rolfsmeier, University of Nebraska,
revealed A. prostrata, an introduced, European annual previously undocu-
mented in Kansas. Kansas specimens all had been mistaken for A. patula
L., which appears to be rare in the Great Plains.

Voucher specimens: KANSAS. Barton Co.: T18S, R12W, sec 9, NW 1/4, SW 1/4, Cheyenne
Bottoms, pool 3, dry, flat, exposed, vegetation sparse, 18 Jul 1985, Brooks 17543 & Laushman.
Edwards Co.: Kinsley, 2 mi E, floodplain of Arkansas River, wet, sandy soil, 5 Oct 1972,
Stephens 63220. Ford Co.: Dodge City, 5 mi E, low bank of Arkansas River, moist, sandy
soil, 18 Sep 1973, Stephens 73993. Gray Co.: Cimarron, 0.5 mi S, bank of Arkansas River,
brushy, weedy, moist, sandy soil, 4 Aug 1971, Stephens 50414. Kearny Co.: Lakin, 4 mi
NE, dry bottom of Lake McKinney, sandy, silty soil, 10 Aug 1975, Stephens 87379.
Berberis thunbergii DC. (Berberidaceae).—Japanese barberry is cultivated
widely as a hedge plant and ornamental. During recent field surveys of the
Ft. Leavenworth Military Reservation in northeast Kansas, widely scattered
plants were encountered 1 in open woods and mesic forests.

Voucher s:s KANSAS. Leavenworth Co.: T8S, R22E, sec 15, SE 1/4, Ft. Leavenworth
Military Recmenon. W-central part, elev 900-1050 ft, mesic, upland, oak-hickory-maple-
basswood forest on slopes SE of Wagner Point, steep draws with few limestone outcrops,

18 May 1995, Freeman 7132; T8S, R22E, sec 23, SE 1/4, Ft. Leavenworth Military Res-
oe SE corner along Corral Creek, E and W along Grant Ave., elev 770-800 ft, open
banks of creek and wooded riparian area immediately W of Grant Ave., 12 Oct 1995, Freeman

Bromus nottowayanus Fernald (Poaceae)—Nottoway brome grows in mesic
forests from southern Quebec south to northern Georgia and east to east-
ern Texas (Pavlick 1995). It may be confused with B. pubescens Muhl. ex
Willd., from which it is distinguished by its sericeous lower leaf sheaths

596 Sipa 18(2

—

and densely pilose collars. A population of B. nottowayanus discovered in
northeast Kansas extends its range westward from Missouri. At the site, it
grows in close proximity with B. pubescens. However, b. nottowayanus is
strictly confined to the narrow floodplain of a small, upland, intermittent
stream in mesic, oak-hickory forest, whereas B. pubescens is much more
widespread, occupying gentle slopes and nde throughout the forest.

/oucher specimen: KANSAS. Leavenworth Co. ce 8S, R22E. sec 11, NW 1/4, Ft.
Leavenworth Military Reservation, central part ENE sf Hancock Hill at Fort de Cavaginal
Picnic Area, elev 860 is ft, mesic to dry-mesic, oak-hickory forest on hills and bluffs
above Missouri River, 21 Jul 1995, Freeman 7448.
Calibrachoa seas (Juss.) D’Arcy (Solanaceae).—Streamside petunia
is a prostrate annual known to occur throughout the southern U.S. an
tropical America (Correll & Johnston 1970; Nee 1993), with historical collections
north to Oregon (Halse 1996). A large population was discovered in cen-
tral Kansas where it occurs in seasonally wet basins. This is the northern-
most locality for the species in the Great Plains. It is possible that this spe-
cies has been overlooked in the Great Plains due to the ephemeral nature
of its habitat.

Voucher specimen: KANSAS. Barton Co.: T18S, RI2W, sec 7, SW 1/4 & sec 8, E 1/2,

Hoisington, 2 mi S, 5 mi W, Cheyenne Bottoms Nature Conservancy Reserve, near Rush

im

ake, elev 1960 ft, barren, dry basins in gr azed, seasonally wet, Pascopyrum-Distichlis grassland,
soil clayey, 17 Jun 1997, Morse 1586 & Loring
Chenopodium pumilio R. Br. (Chenopodiaceae).—Ridged goosefoot is an
annual plant native to Australia and reported over much of the eastern U.S.,
southeastern Oklahoma, much of Texas, and on the West Coast (Gleason
& Cronquist 1991; Hatch et al. 1990; Wilken 1993). A central Kansas collection
extends its range some 270 uses westward from Missouri

Voucher specimen: KANSAS. Reno Co.: 1225S, R7W, sec 10, NE 1/4, iy eooas 9011
N Riverton Rd., common in farm yard, | ie 992, Foster s.

Digitalis lanata Ehrh. (Scrophulariaceae).—Grecian niles is a biennial
or perennial sometimes grown as an ornamental or cultivated for the pro-
duction of digitalis, a drug used as a cardiac stimulant (Moe & Farah 1975).
It is a native of the Balkan Peninsula, Hungary, and Romania. In 1994 it
was reported in southeast Kansas by a landowner in whose scrub-oak woodland
pasture plants were very common and spreading rapidly. This is the only
report of this species naturalizing and becoming noxious in North America.
Since 1995, the Plant Health Division, Kansas Department of Agriculture,
has been attempting to eradicate the plant, a task made difficult by the large
seed bank apparently present at the site.

ee specimens: KANSAS. Wilson Co.: 1285, ves sec 7, SE 1/4, SE 1/4, Fredonia,
6 mi ae 4 mi W, sandy scrub oak pasture, common over ca 20 acres, Mz eh at Vaughn

27 Jun 1994, McGregor 41110 & Russell, 8 Jun | 995, McGregor 411

Dirca spalucian L. (Thymelaeaceae).—Leatherwood grows on on slopes

FREEMAN et al., Plants new to Kansas 597

and along floodplains in mesic, deciduous forests in the eastern U.S. and
southeastern Canada. A thriving population discovered in extreme eastern
Kansas extends the range of this shrub nearly 120 kilometers westward from
central Missouri (Steyermark 1977). The plants first were discovered around
1990 when a site analysis was conducted and land use plans were being
developed for the Overland Park Arboretum and Botanical Gardens, which
now occupies the site.

Voucher specimen: KANSAS. Johnson Co.: THAS, R24E, sec 25, S 1/2, SE 1/4 & sec
26, N 1/2, NE 1/4, jct US 69 & US 169, 6 mi S, 1 mi W, Overland Park Arboretum and
Botanical Gardens, elev 900-1000 ft, mesic, we nikon ie on NE- ee slope S of
Wolf Creek and small limestone glades above steep lin rops, 8 May 1998, Free-
man 10700.

Elaeagnus umbellata Thunb. (Elaeagnaceae).—Autumn olive is an Asian
species planted in the U.S. as an ornamental and for wildlife habitat. Scat-
tered, mature plants occupy clearings and disturbed sites on the Ft. Leavenworth
Military Reservation in northeast Kansas. The first collections from non-
cultivated plants in Missouri were made in the late 1980s, but the species
is believed to be naturalized in most Missouri counties and is a serious pest
(Smith 1997).

Voucher specimen: KANSAS. Leavenworth Co.: T8S, R22E, sec 11, NW 1/4, Ft.

Leavenworth Military Reservation, central part ENE of Hancock Hill at Fort de Cavaginal
Picnic Area, elev 860-1000 ft, mesic to dry-mesic, oak-hickory forest on hills and bluffs
above Missouri River, open, disturbed areas along roads and trails, several dozen scattered
shrubs, 21 Jul 1995, Freeman 7441.
Elodea bifoliata H. St. John (Hydrocharitaceae).—This aquatic macrophyte,
which is native to parts of the Great Plains, Rocky Mountains, and Pacific
Northwest, is added to the Kansas flora based on specimens annotated in
1995 by Robert R. Haynes, University of Alabama. The three specimens
originally were identified as E. nuttallii (Planch.) H. St. John, another member
of the genus that occurs in a few, scattered localities in the state.

Voucher specimens: KANSAS. Cheyenne Co.: St. Francis, 9 mi SW, small seepage feed
pool along Republican River beneath county road bridge, 5 Aug 1980, Brooks 14822; St
Francis, 4 mi SW, pool in South Fork Republican River, 15 Oct 1980, McGregor 32079;
St. Francis, 3.5 mi SW, pool along South Fork Republican River, 16 Aug 1983, McGregor
34761.

Eragrostis pectinacea (Michx.) Nees var. miserrima (E. Fourn.) Reeder
(Poaceae).—This variety (as £. tephrosanthos Schult.) was included in syn-
onymy in the Flora of the Great Plains (Great Plains Flora Association 1991).
However, following Reeder’s (1986) treatment of the species, we have vouchers
for var. miserrima from the following eastern Kansas counties: Allen, Anderson,
Bourbon, Coftey, Douglas, Franklin, Jefferson, Labette, Leavenworth, Linn,
Lyon, Marion, Miami, Morris, Osage, Shawnee, Wilson, and Woodson.
The rangewide distribution of this variety is from Florida and the southern
Gulf Coast west to southeastern Arizona and south through Mexico, Cen-

598 Sipa 18(2)

tral America, and the West Indies (Koch 1974). Variety pectinacea, which
is widespread in the U.S., Mexico, and Central America (Koch 1974), is

common statewide A

Euonymus fortunei (Turcz.) Hand.-Mazz. (Celastraceae).—This climbing
shrub is a frequently planted ornamental in eastern Kansas. Large, well-
established, naturalized populations along the Missouri and Wakarusa Rivers
in eastern Kansas recently have come to our attention. At these sites, F.
fortunei forms dense, spreading carpets in the understory of disturbed floodplain
forests. This species has been seen at numerous other sites in eastern Kan-
sas, most of which have not been documented by vouchers. The varietal
status of these plants has not been determined.

Voucher specimens: Cherokee Co.: 1355, R35E, sec 1, SE 1/4, NE 1/4, Baxter Springs,

E side, Kiwanis Park, elev 200 m, weedy bottomland forest along Spring River, 30 Au
1998, Morse 2723. Douglas Co.: University of Kansas West Campus, woodland west of
Bridwell Lab, undisturbed for over 50 years, 16 Oct 1995, McGregor 41197A, 41197B.
Leavenworth Co.: T8S, R22E, sec 23, SE 1/4, Ft. Leavenworth Military Reservation, SE
corner along Corral Creek, E and W of Grant Ave, open banks of creek and wooded ripar-
ian area immediately W of Grant Ave, 12 Oct 1995, Freeman 7868. Reno Co.: T25S, R8W,
sec 3, NW 1/4, SE 1/4, Arlington, 0.5 mi N, 0.75 mi W, elev 1590 ft, sandy, spring-run
below a spring-fed pond on ne S side of the North Fork Ninnescah River, trunks of larg-
est vines 30—40 cm in diameter, other smaller vines scattered E of pond, 13 Jul 1997, Free-
man 8889.
Eupatorium purpureum L. var. holzingeri (Rydb.) E. Lamont (Asteraceae).—
Lamont (1990; 1995) distinguished two varieties of widespread Eupato-
rium purpureum based on leaf pubescence. Variety purpureum has glabrous
or nearly glabrous abaxial leaf surfaces and occurs throughout most of the
range of the species. In var. holzingeri, the abaxial leaf surface is densely
and persistently puberulent to villose. This variety occurs largely in the Midwest
and eastern Great Plains. An examination of specimens at KANU reveals
both varieties are present in Kansas, with var. purpureum documented only
from Cherokee Co. We have vouchers for var. olzingeri (Rydb.) E. Lamont
from the following 21 counties; Allen, Anderson, Atchison, Bourbon, Cherokee,
Coffey, Crawford, Doniphan, Douglas, Franklin, Jefferson, Johnson, Labette,
Linn, Leavenworth, Miami, Neosho, Osage, Shawnee, Woodson, and
Wyandotte.

Leptochloa uninervia (J. Pres]) Hitchc. & Chase (Poaceae).—Mexican
sprangletop is added to the Kansas flora based on a single specimen anno-
tated in 1990 by Neil Snow, Washington University. This is the only Great
Plains record of this grass, which generally ranges from southern Texas south
into Central and South America (Gould 1975).

Voucher specimen: KANSAS. Rice Co.: T21S, ROW, sec 21, NE 1/4, SW 1/4, Alden,
2.75 mi S, S side of Arkansas River, sandy river floodplain, open, 14 Sep 1983, Brooks
17036.

FREEMAN et al., Plants new to Kansas 599

Lespedeza bicolor Turcz. (Fabaceae).—This perennial legume was intro-
duced from Asia and now is naturalized throughout much of the south-
eastern U.S. (Isely 1990). A small, persistent population found in eastern
Kansas extends the range of the species more than 300 kilometers from
localities in eastern Missouri (Miihlenbach 1979) and northern Arkansas
(Smith 1988).

Voucher specimen: KANSAS. Douglas Co.: T135, R18E, sec 8, N 1/2, SE 1/4, W side
of Clinton Reservoir, Woodridge Public Use Area, elev 1000-1010 ft, weedy, annually
mowed, reseeded tallgrass prairie, scattered wet depressions, 16 Sep 1996, Morse 1077, 9
Oct 1996, Morse 1109.

Loeflingia squarrosa Nutt. subsp. texana (Hook.) Barneby & Twisselm.
(Caryophyllaceae).—This diminutive, native annual grows in sandy soil
from east-central Texas north to north-central Oklahoma, with widely dis-
junct populations in Dawes Co., Nebraska (Barneby & Twisselmann 1970)
and Weston Co., Wyoming (Hartman et al. 1985). Smyth (1898) reported
it from Wichita Co., Kansas, but no voucher has been located. The discov-
ery of a population in southwest Kansas supports its inclusion in the state’s
ora.

Voucher specimen: KANSAS. Clark Co.: T34S, R23W, sec 35, § 1/2, Ashland, 11 mi
S, 0.5 mi W, Dunne Ranch, SE part, elev 1850-1870 ft, nu mixed grass prairie on S
side of the Cimarron River, 28 Apr 1998, Freeman 10588
Lupinus argenteus Pursh var. argenteus iced Sie lupine is a
common species on rocky, mixed grass and shortgrass prairies throughout
the High Plains (Great Plains Flora Association 1977, 1991). A popula-
tion found in northwest Kansas is nearly 200 kilometers from the nearest
locations in western Nebraska and eastern Colorado.

Voucher specimens: KANSAS. Rawlins Co.: T5S, R35W, sec 7, E 1/2 & sec 8, W 1/2,
McDonald, ca 11 mi S, 6 mi E, N-S trending draw on the S side of Beaver Creek, E of the
Dewey Ranch, elev 3200-3300 ft, grazed, upland, shortgrass prairie and rocky slopes be-
low outcrops of the Ogallala Formation, 20 Jun 1990, Freeman 3135, 30 May 1991, Free-
man 3522 e& Brooks
Malus coronaria (L.) Mill. var. coronaria (Rosaceae).—Wild crab apple is
a low shrub or small tree of thickets and woods in the northeastern U.S.
(Gleason & Cronquist 1991). A small, presumably native population was
discovered in extreme eastern Kansas. This species has been documented
in scattered counties in Missouri (Steyermark 1977), including Jackson and
Clay, which are just east of the Kansas locality.

Voucher specimen: KANSAS. Wyandotte Co.: T11S, R25E, sec 34, NW 1/4, Roeland
Park, N side of town, Rosedale Park, elev 850-950 ft, open, oak-hickory forest on pre-
dominantly N-facing slopes S of Turkey Creek, forest slightly to moderately disturbed, 4

y 1998, Freeman 10698.
Malus floribunda Siebold ex Van Houtte (Rosaceae).—This handsome, non-

native, ornamental tree is cultivated widely in eastern Kansas. It is added

600 Sipa 18(2)

to the flora based on a collection from the east-central part of the state.
Voucher specimen: KANSAS. Douglas Co.: University of Kansas West Campus, brushy

woodland SW of greenhouse, area originally an open wooded pasture, undisturbed for 45
years, 7 Apr 1995, Me Gregor 41140.
Paspalum dissectum (L.) L. (Poaceae). — This mat-forming grass grows along
muddy streambanks, ditches, and the margins of ponds in the southeast
U.S. and in Cuba (Gould 1975; Steyermark 1977). Populations in south-

east Kansas extend the range of the species westward 120 kilometers from
the nearest Missouri sites aan liyeee

SA

Voucher specimens: KA oodson Co.: 1268, RI4E, sec 11, Woodson County
State L me on dry lake shore oe common, 13 Oct 1988, McGregor 39793; T25S, RISE,
sec 15 1/4, Yates Center, ca 0.25 mi SW, moist area in drained lake bed, 2 Sep 1993,

ve 41064.
Polygonum cespitosum Blume var. longisetum (Bruijn) Steward
(Polygonaceae).—This introduced, Asian annual has been reported from
several localities along the Missouri River in Iowa and Nebraska (Great Plains
Flora Association 1991). Mitchell and Dean (1978) report the species has
spread rapidly in the U.S. since the 1940s. In recent years, populations 1 in
two northeast Kansas counties have come to our attention. In 1998, it was
observed to be an abundant weed in pots and on the grounds of a plant
nursery in Douglas County, Kansas.
foucher specimens: KANSAS. Douglas Co.: Lawrence, E of Heatherwood Drive, 100

m NW of Heatherwood and 22nd ee disturbed area behind Shannon Plaza Apart-
ments, 23 Oct 1996, Kaderly 15. Leavenworth Co.: T8S, R23E, sec 18, NW 1/4, Ft. Leavenworth
Military Reservation, E-central part, Missouri River floodplain, 1 mi ENE of S end of
Sherman Army Airfield, elev 765 ft, open, silty river bank and mesic, floodplain forest, 3
Oct 1995, Freeman 7779; T8S, R22E, sec 23, SE 1/4, Ft. Leavenworth Military Reserva-
tion, SE corner along Corral Creek, E and W of Grant Avenue, open banks of creek and
wooded riparian area immediately W of Grant Avenue, 12 Oct 1995, Freeman 7864.
Rhamnus cathartica L. (Rhamnaceae)—Common buckthorn is a Euro-
pean shrub or small tree cultivated in the U.S. as a hedge plant. It has be-
come naturalized and occasionally weedy in parts of the eastern U.S. and
southern Canada (Elias 1980; Smith 1997). Scattered, small trees were discovered
in a disturbed forest in extreme eastern Kansas.

Voucher specimen: KANSAS. Wyandotte Co.: T11S, R25E, sec 34, NW 1/4, Roeland

Park, N side of town, Rosedale Park, elev 850-950 ft, open, oak-hickory forest on pre-
dominantly N-facing slopes S of Turkey Creek, 4 May 1998, Freeman 10677.
Schoenoplectus californicus (C.A. Mey.) Sojak (Cyperaceae).—A single
specimen of California bulrush was discovered among KANU specimens
of Schoenoplectus in 1993 by S. Galen Smith, University of Wisconsin-
Whitewater, while he Was annotating specimens for the Flora of Nor th America
project. As far as we are aware, this is the only record of this taxon in the
Great Plains. It differs from other members of the Schoenoplectus lacustris
complex (S. tabernaemontani (C.C. Gmel.) Palla, S. acutus (Muhl. ex Bigelow)

FREEMAN et al., Plants new to Kansas 601

A. Love & D. Live, and S. heterochaetus (Chase) Sojak) in having bluntly
trigonous culms near the inflorescence, leaf blades lacking or up to 2 mm
long, strap-like floral bristles, and smooth floral scales. In the U.S., Schoenoplectus
californicus is distributed discontinuously from the southern Atlantic Coastal
Plain west to California.

Voucher specimen: KANSAS. Barber Co.: 2 mi SE of Sun City, edge of creek bank, 4
Aug 1959, McGregor 14712.
Shepherdia argentea (Pursh) Nutt. (Elaeagnaceae).—Smyth (1892), Hitchcock
(1899), and Gates (1940) listed buffaloberry in Kansas, but we are unable
to locate a voucher substantiating the historical occurrence of this species
in the state. This is a common, widespread shrub in the northern half of
the Great Plains. A small population in northwest Kansas confirms its presence
in Kansas and extends the species’ range south from Nebraska roughly 40
kilometers.

oucher specimen: KANSAS. Rawlins Co.: T4S, R36W, sec 32, ca 8.5 mi S, 5.5 mi E

of McDonald, SE-trending en that joins eas Creek to the S, elev 3100-3160 ft,
grazed, upland, sl f the Ogallala Formation,
resthicted toaca 50 yd eiretoh of the N bank fF Beaver Creek, 19 Sep 1990, Freeman 3248
e Brooks

Sida chombifolia L. (Malvaceae).—No other reports of this pantropical annual,
which occurs sporadically in ruderal sites in the eastern U.S. (Fryxell 1985;
Gleason & Cronquist 1991), are known for the Great Plains. When it was
collected, the land owner reported the population had been present at least
10 years. It is not known if this population persists.

Voucher specimen: KANSAS. Shawnee Co.: Topeka, backyard of home at 1916 SW
Atwood, lawn weed at edge of garden, known from the site for at least 10 years, 5 Oct
1992, Coleman s.n.

Stellaria pallida (Dumort.) Crép. (Caryophyllaceae).—Lesser chickweed
first was reported for North America by Morton (1972), who collected it
in North Carolina. Rabeler (1997) reported it for Texas and noted that it

had been found in nearby states, including Arkansas and Tennessee.
Our field and herbarium studies reveal this AeA weed of lawns and dis-
turbed areas occurs in 84 Kansas counties. In the past it has been misidentified

as the ubiquitous Stellaria media (L.) Vill., common chickweed. Rabeler
(1997) considered S. pallida to bea probable recent introduction, but specimens
indicate it has been in Kansas at least 67 years. The oldest collections we
have seen are deposited at the Kansas State University Herbarium (KSC):
a 1931 collection from Lyon Co. and a 1946 collection from Cherokee Co.
Tridens xoklahomensis (Feath.) Feath. (Poaceae) —Oklahoma tridens is
a naturally occurring, sterile hybrid between 7ridens flavus (L.) Hitchc. and
T. strictus (Nutt.) Nash (Crooks & Kucera 1973). It currently is known

rom two sites in southeast Kansas, and additional field studies likely will

602 Sipa 18(2)
reveal other eee where the parent species are sympatric.

Voucher specimens: KANSAS. Labette Co.: Parsons, near the arboretum, native-like
area, 2 Oct 1995, Holland a Neosho Co.: St. Paul, 1 mi N, open pasture, growing
with 7ridens strictus, 19 Oct 1995, Holland 8583.

Trifolium incarnatum L. (Fabaceae).—Records of crimson clover in the
Great Plains are based on infrequent escapes from plantings (Great Plains
Flora Association 1991). This showy, European legume was found at a single
locality in east-central Kansas where the land owner reports it has persisted
for at least four years.

Voucher specimen: KANSAS. D ea ee T128, RIE, see 21, E 1/2, 5'W 1/4, jet
KS 10 & US 40 (W of Lawrence), 1. a , 1.5 mi E, elev 890-960 ft, disturbed, oak-
hickory woodland on N-S ridge between two unnamed tributaries to Baldwin Creek, mowed
path on E side of ridge, 5 May 1 998, Freeman 10699
Veronica biloba L. Geeta tence ieedehs speedwell, an annual
introduced from western Asia, has been reported in the Great Plains from
ruderal sites along the Front Range of the Rocky Mountains (Great Plains
Flora Association 1991) and from two recent collections in Nebraska. It is
ea to the Kansas list based on a collection in north-central Kansas.

ucher specimen: SAS. Ottawa Co: T11S, R4W, sec 1, SE 1/4, Minneapolis, S

a. es: town in Markley Grove Park, elev 1240 ft, wooded floodplain on S side of Solomon

River, woods eae by Quercus macrocarpa, understory mostly mowed, 6 May 1997,
Freeman 867°

Veronica ee L. (Scrophulariaceae).—Ivy-leaf speedwell is a widely
introduced, European species known from three previous collections in the
Great Plains: two in Nebraska and one in South Dakota (Great Plains Flora
Association 1977, 1991). Populations discovered in Lawrence, Kansas, in
1993 and 1995 are persisting.

Voucher specimens: KANSAS. Douglas Co.: T12S, RI9E, sec 31, Lawrence, NW of
the University of Kansas campus, W side of Emery Rd, ca 30 ft S of jct with Sigma Nu
oe Sear open, ile weedy, E-facing slope, 17 Apr 1993, Freeman 261. 4

993, ee 47783 T13S, R20E, sec 6, NW 1/4, NW 1/4, Lawrence, alley on W
aes f Tennessee just S of corner of 16th and Tennessee, oe ground cover along
a and behind soins at 1627 Tennessee, 24 Apr 1995, 2uF: ene Lake, lawn
of headquarters office, 20 Apr 1997, oe yy 292. Franklin Co.: In Ottawa, lawn of
city park, 19 Apr 1997, McGregor 412:

Veronica persica Potr. ee ee speedwell is another
widely introduced, European annual known from a few sites in the Great
Plains. It was inadvertently omitted from the Flora of the Great Plains (Great
Plains Flora Association 1991), although it has been documented in lowa,
Nebraska, South Dakota, and Wyoming. A 1975 collection from western
Kansas went unnoticed until recently, and it is not known if the species
still occurs at the site.

Voucher specimen: KANSAS, Scott Co.: Scott City, park at south edge of city, scat-
tered in lawn, 26 Jul 1975, McGregor 27516.

FREEMAN et al., Plants new to Kansas 603
ACKNOWLEDGMENTS
We wish to thank R. Laurie Robbins (KSTC) and Robert B. Kaul (NEB)

for their suggestions and comments on the manuscript. Support for some
field travel was provided to CCF and CAM by the Kansas Biological Sur-

vey, University of Kansas.

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AL-SHEHBAZ, I.A. and V. Bares. ei Armoracia lacustris (Brassicaceae), the correct name
for the North American lake cress. J]. Arnold Arbor. 68:357-—359.

Barnesy, R.C. and E.C. jon 1970. Notes on Loeflingia (Caryophyllaceae). Madrono
20:398-407.

CorreLl, D.S. and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Yexas
Research Foundation, Renner.

Crooks, P and C.L. Kucera. 1973. Tiridens x ok aaa flavus x T. strictus), an inter-
specific sterile hybrid in the Eragrosteae (Gramineae). Amer. J. Bot. 60:262—267.

Euias, T.S. 1980. The complete trees of North ens field guide and natural history.
Van Nostrand Reinhold Co., New York, NY.

FryxeLL, PA. 1985. Sida sidarum - V. The North and Central American species of Sida.
Sida 11:62-91

Gates, EC. 1940. Flora of Kansas. Agric. Exper. Sta., Kansas State Coll. Agric., Manhat-

tan.

GLEASON, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United
States and adjacent Canada. New York Botanical Garden, Bronx.

GOouLb, F.W. 1975. The grasses of Texas. Texas A&M University Press, College Station.

GREAT PLAINS FLORA ASSOCIATION. 1977. Atlas of the flora of the Great Plains. Iowa State
University Press, Ames

GreaT PLAINS FLORA ASSOCIATION. 1991 (second printing). Flora of the Great Plains. Uni-
versity Press of Kansas, Lawrenc
Hatse, R. 1996. Noteworthy dallections: Oregon. Madrofio 43:98.

Hartman, R.L., B.E. NeLson, and K.H. DurHoLm. 1985. Noteworthy collections: Wyo-
ming. Madrofio 32:125-128.

Haren, S.L., K.N. GANDHI and L.E. Brown. 1990. Checklist of the vascular plants of
Texas. Texas Agric. Exp. Sta. MP-1655. Texas A & M Univ., College Station.

HENRICKSON, J. 1993. Solanaceae. In: J.C. Hickman, ed. The Jepson onal Higher plants
of California. University of California Press, Berkeley.

Hitchcock, A.S. 1899. Flora of Kansas; a series of maps illustrating the distribution of
flowering plants by counties. Kansas State Agric. Coll., Manhattan.

IseLy, D. 1990. Vascular flora of the southeastern United States. vol. 3, pt. 2: Leguminosae
(Fabaceae). ey of North C arolina Press, Chapel Hill.

Kocn, $.D. 1974. The Eragrostis pectinacea-pilosa complex in North and Central America
(Gramineae: Eragrostoideae). Illinois Bical. Monogr. 48:1—74.

LAMONT, E.E. 1990. A new combination in Eupatorium section Verticillata (Asteraceae).
Phytologia 69:467—468.

LAMONT, E.E. 1995. gas of Eupatorium section Verticillata (Asteraceae). Mem. New
York Bot. Gard. 72:1

MITCHELL, R.S. and J.K. AN. 1978. Polygonaceae (buckwheat family) of New York state.
New York State Mus. Bull. 431:1-79.

Mok, G.K. and ALE. Faran. 1975. Digitalis and allied cardiac glycosides. In: L.S. Goodman
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Co., Inc. New York, NY.

Morton, J.K. 1972. On th u f Stellaria pallida in North America. Bull. Torrey
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MUHLENBACH, V. 1979. Contributions to the synanthropic (adventive) flora of the rail-
roads in St. Louis, Missouri, USA. Ann. Missouri Bot. Gard. 66:1 :

Neg, M. 1993. Solanaceae. In: J.C. Hickman, ed. The Jepson er Higher plants of
California. University of California Press, Berkeley.

Pavuck, L. 1995. Bromus L. of North America. Royal British Columbia Mus., Victoria,

BC.

RABELER, R.K. 1997. Cerastium pumilum and Stellaria pallida new for Texas. Sida 17:843-
845.

RE - a R. 1986. Another look at Eragrostis tephrosanthos (Gramineae). Phytologia 60:153—

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NEW AND NOTEWORTHY ANGIOSPERM
RECORDS FOR IOWA

CRAIG C. FREEMAN

R.L. McGregor Herbarium, Division of Botany
Natural History Museum and Biodiversity Research Center
University of Kansas, 2045 Constant Ave.
Lawrence, KS 66047-3729, U.S.A.
c-freeman@ukans.edu

Recent field work in Iowa has resulted in the discovery of three angiosperms
previously unknown in the state and populations of nine other angiosperms
that are rare. Eight of the 12 species are considered native in Iowa. Most
populations were discovered during field surveys that are part of an effort
to complete a comprehensive flora of Bremer County, located in northeast
Iowa. One species excluded from the Iowa flora by recent workers should
be reinstated based on a 1998 collection in the state and an older herbarium
specimen at the R.L. McGregor Herbarium (KANU), University of
Kansas.
NEW STATE RECORDS

Eleocharis intermedia (Muhl.) Schult. (Cyperaceae)—A number of au-
thors have included Iowa within the range of this cespitose spike-rush (Rydberg
1932; Svenson 1957; Mohlenbrock 1976; Gleason & Cronquist 1991); however,
it was not listed for the state by Eilers and Roosa (1994). A small popula-
tion discovered along the Cedar River in 1997 supports its inclusion in the
state’s flora.

Voucher specimen: IOWA. Bremer Co.: T92N, R14W, sec 22, SW 1/4, SW 1/4 & sec
21, SE 1/4, SE 1/4, Waverly, 0.5 mi W, 1.5 mi N, Cedar Bend Conservation Park, E end,
drying mud flats along the Cedar River, 7 Oct 1997, Freeman 10198 (ISC, KANU),
Rumex stenophyllus Ledeb. (Polygonaceae).—Eilers and Roosa (1994) excluded
this introduced species from their checklist, stating that reports of the taxon
in Iowa were unsubstantiated. It was mapped in Harrison County, Iowa,
in the Adlas of the Flora of the Great Plains (Great Plains Flora Association
1977) based on a voucher at KANU, and it recently was collected at a sec-
ond locality in the state.

Voucher specimens: IOWA. Bremer Co.: T9IN, R14W, sec 2, SE 1/4 NW 1/4, Waverly,
E side of town, open, disturbed railroad right-of-way just S of Bremer Avenue, 18 Jul 1998,
Freeman 11090 (ISC, KANU). Harrison Co.: 3 mi SE of Modale, margin of farm pond,
exposed, weedy, wet, 17 Aug 1974, Brooks 8173, McGregor & Stephens (KANU).

SIDA 18(2): 605-609. 1998

606 Sipa 18(2)

Spergularia marina (L.) Griseb. (Caryophyllaceae).—This diminutive,

introduced, annual weed occurs sporadically in eae North America and
more commonly along the coasts (Rossbach 1940). A small population was
discovered in 1991 in a disturbed, grassy site at the edge of a sidewalk ina
rest area along Interstate 35. More plants were discovered in a grassy me-
dian in the parking lot of the same rest area in 1995. Annual visits reveal
this population i is persisting.

Voucher specimens: IOWA. Clarke Co.: roadside rest area along I-35, ca 2 mi W & 2
mi S of Osceola, W side of interstate, 2 Sep 1991, Freeman er Freeman 4124 (KANU); 5
Sep 1995, Freeman ce» Freeman 7766 (ISC, KANU)

NOTEWORTHY RECORDS

Astragalus distortus Torr. & A. Gray var. distortus (Fabaceae),—Bent milk-
vetch is documented in 11 southeastern and eastern Iowa counties; how-
ever, extant populations occur in only six of these (Roosa et al. 1989). The
species typically is associated with sand prairies (Eilers & Roosa 1994) and
is a special concern plant (lowa Administrative Code 571 1994). A small,
degraded, sandy prairie remnant in a cemetery north of Janesville supports
roughly two-dozen plants of this species. Despite the small size of this population,
many individuals produced fruits in the spring of 1998. This is the first
report of the species from the coun

Voucher specimen: IOWA. Bremer Co.: T91N, R14W, sec 35, NE 1/4, NE 1/4, NE 1/

4, Janesville, 0.5 mi N, Oakville Cemetery, eae sandy, upland cemetery E of the Ce-
dae River, 24 May 1998, Freeman 10757 (ISC, NU).
Besseya bullii (Eaton) Rydb. ee Nee eee kittentail is a
Midwest endemic found on dry prairie ridges, in woodlands, and along
sandy river terraces (Roosa et al. 1989). It has been documented in more
than a dozen eastern and north-central Iowa counties, including Bremer
County. It is a threatened species in the state (lowa Administrative Code
571 1994). A new population of fewer than 20 individuals was discovered
in Bremer County in 1997, and because the population is small, a voucher
specimen was not collected. Instead, a photograph was taken and serves as
the voucher for this population.

Voucher specimen: IOWA. Bremer Co.: T91N, R14W, sec 21, W 1/2, Janesville, 2 mi
N, 2.5 mi W, Shell Rock Greenway, Shell Rock Access, mesic floodplain forest along a
abner to the Shell Rock River and small area of remnant tallgrass ae along an old

U)

road right-of-way, 3 Jun 1997, Freeman s.n. [photographic vouchers] (K

Carex conoidea Schkuhr ex Willd. (Cyperaceae).—Field sedge is a special
concern plant in lowa (lowa Administrative Code 571 1994). It is docu-
mented from Black Hawk, Bremer, Fayette, Linn, and Scott counties in
eastern Iowa (Mark Leoschke, pers. comm.; Roosa et al. 1989). A second
Bremer County population recently was discovered in a mesic prairie growing
with C. bicknellii Britton, C. cristatella Britton, C. sartwellii Dewey, and C.
stricta Lam.

FREEMAN, Angiosperm records for lowa 607

ae specimen: IOWA. Bremer Co.: T92N, R13W, sec 29, NE 1/4, NE 1/4, NE 1/
4, Bremer, 1.1 mi S, elev 1000 ft, small, mesic, upland, tallgrass prairie remnant alon
intermittent stream on W side of blacktop, soil rich and loamy, infrequent, 19 Jun 1998,

Freeman 11055 (ISC, KANU).
Carex siccata Willd. (Cyperaceae).—The only previous record of this sedge
in Iowa is a 1936 collection from Clay County (Gilly 1946; Roosa et al.
1989), where it was reported as C. foenea Willd. However, the name Carex
foenea has been misapplied in much of the literature (Cronquist et al. 1977;
Gleason & Cronquist 1991). While I have not seen the voucher cited by
Gilly (1946), his key and description match C. siccata rather than C. foenea.
‘True C. foenea is a member of section Ovales and occurs north of Iowa.
Carex siccata, a member of section Arenariae, is widely distributed in the
northern U.S. and southern Canada (Cronquist et al. 1977). It is listed (as
C. foenea) as presumed extirpated by Roosa et al. (1989) and is a special
concern plant (Iowa Administrative Code 571 1994). A small population
was discovered in a sand prairie remnant along the Wapsipinicon River,
where Stipa spartea Trin. is the dominant species. Common species at the
site are Dichanthelium depauperatum (Muhl.) Gould, Euphorbia corollata
L., Plantago patagonica Jacq. var. patagonica, Poa pratensis L., Lespedeza capitata
Michx., Rumex acetosella L., and Vulpia octoflora (Walter) Rydb

Voucher specimens: IOWA. Bremer Co.: T92N, R12W, sec 2, E 1/2, Tripoli, 1.5 mi E,
Sweet Marsh Wildlife Management Area, S end, small, sandy prairie remnant on the E
side of the Wapsipinicon River 4 Jun 1997, Freeman 9018 (KANU); 16 Jun 1997, Free-
man 9234 (KANU, ISC).

Floerkea proserpinacoides Willd. (Limnanthaceae).—This weak, forest annual
is reported only from Dubuque County (Eilers & Roosa 1994). It is a spe-
cies of special concern (Iowa Administrative Code 571 1994). A large colony
was discovered in a narrow, forested, creek floodplain southeast of Waverly.

Voucher specimen: IOWA. Bremer Co.: T91N, R13W, sec 16, NW 1/4, SW 1/4, Waverly,
3 mi E, 1.5 mi S, rich, mesic, upland, maple-basswood forest and oak-hickory forest on
slopes along Baskins Creek and its tributaries, large colony along floodplain of creek, 24
May 1997, Freeman 8947 (ISC, KANU)

Geum vernum (Raf.) Torr. & A. Gray (Rosaceae).—Spring avens is reported
from five lowa counties, mostly in the southern third of the state (Eilers &
Roosa 1994). It is a species of special concern (lowa Administrative Code
571 1994). Scattered plants were discovered in northeast Bremer County
in 1997, the first record for the county.

Voucher specimen: IOWA. Bremer Co.: T93N, R11W, sec 13, N 1/2, NE 1/4, Sumner,
1 mi N. 0.5 mi E, North Woods County Park, mesic, maple-basswood-hickory forest E of
the Little Wapsipinicon River and clearings in the forest, infrequent, 4 Jun 1997, Freeman
9049 (ISC, KANU),.
Platanthera flava (L.) Lindl. var. herbiola (R. Br.) Luer (Orchidaceae),—
This orchid has been reported in low, mesic prairies and sedge meadows in

608 Stipa 18(2)

eight counties scattered in the eastern third of the state (Roosa et al. 1989),
including Bremer County. It is endangered in Iowa (lowa Administrative
Code 571 1994). Two populations were discovered in the Wapsipinicon
River drainage in Bremer County. One population of less than a dozen individuals
(Freeman 9685) was found ina formerly pastured, sedge meadow. This appears
to be the first record of the species at this site. The second population (Freeman
11020) is much larger, comprising more than 100 individuals scattered in
a 2-acre area of mesic tallgrass prairie on the north end of the Sweet Marsh
Wildlife Management Area. This population first was discovered in 1993
by Mark Leoschke of the lowa Department of Natural Resources (John Pearson,
pers. comm.).

Voucher specimens: IOWA. Bremer Co.: T92N, R11 W, sec 20, NW 1/4 & sec 19, NE
1/4, Tripoli, 2.5 mi S, 4 mi E, Wapsi Greenbelt, Buhr Access, open sedge meadow S of
Bremer Co. C33 and E of the ee ares River, 7 Aug 1997, Freeman 9685 (KANU);
T92N, R12W, sec 27, SE 1/4, SE 1/4, SE 1/4, Tripoli, 1 mi N, 1 mi E, Sweet Marsh
Wildlife Management Area, N s side, mesic to wet-mesic prairie W of the East Fork Wapsipinicon
River, 18 Jun 1998, Freeman 11020 USC, KANU).

Platanthera psycodes (L.) Lindl. (Orchidaceae).—Purple fringed orchid
has been documented in 10 counties in northeast and east-central Iowa
where it typically grow in mesic prairies, sedge meadows, and fens (Roosa
et al. 1989). Nekola (1990) reported populations from two sites in Bremer
County. Recently, one additional site for this threatened species (Iowa
Administrative Code 571 1994) was discovered in the same county. A pho-
tographic voucher was taken due to the small size of this population.
Voucher specimen: IOWA. Bremer Co.: T92N, R11W, sec 20, NW 1/4 & sec 19, NE
1/4, Tripoli, 2.5 mi S, 4 mi E, Wapsi Greenbelt, Buhr Access, open sedge meadow S of
Bremer Co. C33 and E of the Wapsipinicon River, 7 Aug 1997, Freeman 8033-8036 | pho-
tographic vouchers] (KANU)
Potentilla intermedia L. (Rosaceae).—This European native previously was
reported only from Black Hawk County, Iowa (Eilers & Roosa 1994). A
small population was discovered in 1997 in adjacent Bremer County.
Voucher specimen: IOWA. Bremer Co.: 191N, ae sec 2, N 1/2, SE 1/4, Waverly,
SE side of town along the Cedar River in Brookwood Park, mesic floodplain forest and
clearing along the river, infrequent along gravelly trail, 16 Jun 1997, Freeman 9205 (ISC,
NU).

ACKNOWLEDGMENTS
Deborah Lewis (ISC) and Ronald L. McGregor (KANU) provided helpful
comments on the manuscript, for which I am grateful. The members of
the Bremer County Conservation Board are acknowledged for their con-
tinuing support of my field work in northeast Iowa.

REFERENCES
CRONQUIST, A., AJH. HOLMGREN, N.H. Houmaren, J.L. REVEAL and PK. HOLMGREN. 1977.

FREEMAN, Angiosperm records for lowa 609

Intermountain flora: vascular plants of the Intermountain west, U.S.A. Vol. 6; the
monocotyledons. Columbia Univ. Press. New York, N
Eiters, L.J. and D.M. Roosa. 1994. The vascular plants of Iowa. Univ. Iowa Press. lowa

ity.
Guty, C.L. 1946. The Cyperaceae of Iowa. Iowa State Coll. J. Sci. 21:55-151.
GLEASON, H.A. and A. Cronquist. 1991. Manual of the vascular plants of the north-east-
ern United States and adjacent Canada. New York Bot. Gard. Bronx.
GREAT PLAINS FLoRA ASSOCIATION. 1977. Atlas of the flora of the Great Plains. lowa State
. Press. Ames, I
Iowa ADMINISTRATIVE CODE 571. 1994. Chapter 77: endangered and threatened plant and
animal species. lowa Dept. Nat. Resources. Des Moines.
MOHLENBROCK, R.H. 1976. The illustrated flora of Hlinois. Sedges: Cyperus to Scleria.Southern
Illinois Univ. Press. Carbondale.
NEKOLA, J.C. 1990. Rare Iowa plant notes from the R.V. Drexler Herbarium. J. lowa Acad.
Sci. 97: 55-73.
Roosa, D.M., M.J. LEoscHKE, and L.J. Erers. 1989. Distribution "a lowas endangered
and threatened vascular plants. lowa Dept. Nat. Resources. Des M
RossBacu, R.P. 1940. Beene in North and South America. Rilo iets 40: 57-83, 105—
143, 158-193, 203-213.
Rypperc, P.A. 1932. Flora of the prairies and plains of central North America. New York
. Gard. Bronx.
SvENSON, H.K. 1957. Eleocharis. N. Amer. Fl. 18:509-540.

610 Sipa 18(2)

BOOK NOTICE

Josrr J. Haba. 1996. The genus Gentiana by Josef J. Halda, Illustrated
y Jarmila Haldova. (ISBN 80-901846-6-9, hbk). Sen, Dobre. Available
through: Harvy Wrightman, Rr#3, Kerwood Ontario, NOM 2B0,
CANADA, Josef J. Halda, PO Box 110, Hradec Krdlové 2, 501 01,
CZECH REPUBLIC. US$60.00 (hbk), $US50.00 (pbk) + US$10
p&h. 241 pp, 300 colored paintings on 32 plates.

“The Genus Gentiana is the first full account of all the species of Gentiana written as a
single book. It is a major taxonomic revision of the whole genus, with an emphasis on
species. A definitive reference work, combining the information needed by botanical, hor-
ticultural, nursery, specialist, and general gardening audienc
“Profusely illustrated by Jarmila Haldova, an exquisite ee appears on every page.
Finally you can see and learn about such elusive plants as Hebe-looking Gentiana pac hyphylla
or the most ornamental species as G. amplicrater, G. grandiflora, G. sedifolia, G. delavayi,
G. szechenyi or G. boissierti which Halda has introducted to cu ultivation in recent years.’
The contents of the book include: Introduction; Morphology; Cultivation; Propaga-
tion; Key to the subgenera; Synopsis of the Genus Gentiana; Sugenus Gentiana; Subge-
nus Stylophora, Subgenus Otophora; Subgenus Pneumonanthe; Subgenus Newberryi; Sub-
genus Ciminalis; Subgenus Cruciata; oo Phyllocalyx; Subgenus Frigida; Subgenus
Monopodiae, Subgenus /someria, Subgenus Microsperma; Subgenus Stenogyne; Subgenus
sani aiienis Subgenus Chondr oe Ca hybrids and cultivars; Related genera;
Speci inally described as Gentians later transferred to the related g genera; Referen ices;

ye a

SIpA 18(2): 610. 1998

LUZIOLA PERUVIANA (POACEAE: ORYZEAE)
PREVIOUSLY UNREPORTED FROM TEXAS AND
A KEY TO TEXAS SPECIES

STEPHAN L. HATCH
S.M. Tracy Herbarium (TAES)
Department of Rangeland Ecology and Management
Texas A&M University, College Station
DX 784 922 126, USA:
s-hatch@tamu.edu

DAVID J. ROSEN

US Army Corps of Engineers
New Orleans District, PO. Box 60267
New Orleans, LA 70160, U.S.A.

JAMES A. THOMAS

Texas Parks and Wildlife Department
Upper Coast Wetland Ecosystem Project
1201 Childers Ra., Orange, TX 77630, U.S.A.

JOHN E. DAWSON III

S.M. Tracy Herbarium (TAES)
Department of Rangeland Ecology and Management
exas AGM University
College Station, TX 77843-2126, U.S.A.

1]

Luziola peruviana Gmel. was collected June 28, 1997, while ing wetland
plant specimens at the J.D. Murphree Wildlife Management Area south-
west of Port Arthur, Texas. Luziola peruviana was identified using Chase
(1951), Allen (1975), and Terrell and Robinson (1974) and verified from
specimens in the S.M. Tracy Herbarium (TAES).

Luziola peruviana has previously been reported to occur in western Florida
(Chase 1951) but specimens were not examined by Hall (1978). Allen (1975)
reported the species from Orleans and Plaquemine Parishes in Louisiana.
A search of Texas floras and checklists does not render a report for this spe-
cies (Correll & Johnston 1970; Gould 1975; Hatch et al. 1990. Jones et al.
LO 7)

Stipa 18(2): 611-614. 1998

612 Sipa 18(2)

A KEY TO LUZIOLA IN TEXAS

—_

. Pistillate inflorescences a raceme or solitary spikelet, axillary; staminate

inflorescences a raceme or panicle, terminal; culms 30-100 cm long,

slender, ics eeodiae leaf blades 2-4 cm long, 2—4(—5) mm er pe
at, linear to lanceolate L. fluitans

—

fle
. Pistillate inflorescences a panicle, axillary, diffuse; staminate inflorescences
a panicle, terminal; culms 21—35 cm long, slender, ascending; leaf blades

10-12 cm long, 2.5-3 mm wide, flat, linear L. peruviana
ame eo (Michx. ) Terrell & H. Robinson, Bull. Torrey Bot. Club
1:235—245. 1974.
Fora a. see Gould (1975) under the synonym Hydrochloa caroliniensis
Beauv.

Luziola peruviana Gmel. Syst. Nat. 2:637. 1791. (Fig. 1).

Plants monoecious, stoloniferous, aquatic. Culms 21—32(—35) cm long,
slender, ascending; nodes often with a dark purple ring below. Leaves cauline;
sheaths open, tessellate; ligules 5—11(—-12) mm long, membranous, lacer-
ate with 3-several hyaline, acuminate divisions; blades (those subtending
pistillate inflorescences) 10—12 cm long, 2.5—3 mm wide, linear, flat, gla-
brous, margins serrulate. Spikelets unisexual, florets disarticulating at the
base; glumes absent. Staminate inflorescences a panicle, terminal, narrow,
exserted above leaves; staminate spikelets 5.3-6.5(—7) mm long, hyaline;
lemmas 7-veined, erose; paleas 5-veined, sub-equal to lemmas; stamens 6;
anthers 3—3.2 mm long, basifixed, yellow, visible through floret. Pistillate
inflorescences a panicle, axillary, not exserted above the leaves, diffuse. Pis-
tillate spikelets (1.8—)2.1—2.5 mm long, glabrous, ovoid, acuminate; lem-
mas 7-veined; paleas 5-veined, sub-equal to lemmas, loosely enclosing mature
caryopses.

The species L. peruviana was flowering when collected in 0.6—0.75 m of
water. I'he water temperature at the site was 28°C and the salinity 0.2 ppt
(fresh water). The associated species included Panicum repens L., Paspalum
distichum L., Limnobium spongia (Bosc.) Steud., Hydrocotyle umbellata L.,
Alternanthera philoxeroides (Mart.) Griseb., Ludwigia peploides H.B.K., and
Ludwigia uruguayensis (Camb.) Hara.

Distribution.—In Texas, along coastal waterways of Jefferson County.

Sp { 7 Jefferson Co.: J.D. Murphree Wildlife Management ae
Big il oe 28 Jun 1997, D. Rosen, S. Hatch, J. Thomas, ]. Sutherlin 364 (TAES);
Hatch, D. en, J. Thomas 6746 (BRIT, TAES); J.D. Murphree Wildlife a nee
Area, Big H oT ne 3.5 mi S of the convergence with Taylor's Bayou, between compart-
ments 4 & 5, 30 Jun 1997, Thomas 73 (TAES), associated with Panicum repens, Paspalum
distichum, and Hydrocotyle umbellata.

Hatcu et al., Luziola peruviana in Texas 613

lem

Fic. 1. Luziola peruviana. A. Plant. B. Pistillate spikelet. C. Staminate spikelet. D. Stami-
nate inflorescence. E. Ligule. F. Pistillate inflorescence. (A-D and F, Chase 1951; E. J.E.
Dawson III).

614 Sipa 18(2)

ACKNOWLEDGMENTS

Financial support provided in part by Texas Higher Education Coordi-
nating Board - Advanced Research Program and the Texas Agricultural
Experiment Station.

REFERENCES

ALLEN, C.M. 1 Grasses of Louisiana. Univ. of Southwestern Louisiana. Lafayette, LA.

Cuase, A. 1951. Hitchcock’s manual of the grasses of the United States, 2nd. ed. U.S.D.A.
Misc. Publ. os 200. U.S. Government Printing Office, Washington, DC.

Corre.i, D.S. and M.C. JOHNSTON. 1970. Mi anual of the vascular plants of Texas. Texas
Research Foundation, Renner, TX.

GouLb, E.W. 1975. The grasses of Texas. Texas A&M Univ. Press, College Station, TX.

Hat, D.W. 1978. The grasses of Florida. University of Florida, Ph.D. Dissertation, Gainsville.

Haren, $.L., K.N. Ganpti, and L.E. Brown. 1990. C ie of the vascular plants of
‘Tex Vass Agric. Exp. Sta. Bull. MP-1655, College Station.

ne. 5 ‘S. eee es Wiper, and PRM. Montcomery. 1997. Vassuler plants of Texas. Univer-
sity of Texas Press, Austin.

Terrewt, E.E., and H. Ropinson. 1974. Luziolinae, a new subtribe of oryzoid grasses. Bull.
Torrey Bot. Club 101:235-245.

oh

LACTUCA SALIGNA (ASTERACEAE),
A LETTUCE NEW FOR TEXAS

ROBERT J. OKENNON

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A., bokennon @brit.org
GEORGE M. DIGGS, JR.
Department of Biology
Austin College
Sherman, TX 75090 U.S.A.
and Botanical Research Institute of Texas, gdiggs@austinc.edu

BARNEY L. LIPSCOMB

Botanical Research Institute of Texas
D Pecan Street

50
Fort Worth, TX 76102-4060, U.S.A., barney@brit.org

Lactuca saligna L., willow-leaf lettuce, is a European native (Ferdkova 1976)
naturalized in a variety of localities in the United States. It is usually de-
scribed as a weed inhabiting roadsides, disturbed places, and waste areas.
The species is widely known from the eastern U.S., with localities reported
from near the east coast (North Carolina, South Carolina, Virginia) west
to Arkansas, Michigan, Missouri, Ohio, Tennessee, and West Virginia (Robinson
& Fernald 1908; Britton & Brown 1913; Fernald 1950; Gleason 1952;
Gleason & Cronquist 1963, 1991; Steyermark 1963; Radford et al. 1968;
Strausbaugh & Core 1978; Cronquist 1980; Smith 1988). Its spread has
apparently occurred mainly during this century because it was reported by
neither Watson and Coulter (1889), Britton and Brown (1898), nor Small
(1903, 1913, 1933). Further, Strausbaugh and Core (1978) indicated that
the species “. .. was not reported by Millspaugh [for West Virginia] in 1913,
which may be an indication that it has spread widely since that time.” It
was, however, reported early in the century for Ohio by Robinson and Fernald
(1908). A relatively late spread to the west is supported by Steyermark’s
(1963) statement that the species was first known from Missouri in 1932.
In addition to the eastern United States, Lactuca saligna has been reported
from the Great Plains (lowa, Kansas, Nebraska, South Dakota—Great Plains
Flora Association 1977; Barkley 1986) and California (Stebbins 1993). It

Sipa 18(2): 615-619. 1998

616 Sipa 18(2)

SS

Fic. 1. Habit of Lactuca saligna (Britton & Brown 1913

is also known from Oklahoma (Taylor & Taylor 1994), with forma ruppiana
(Wallr.) Beck [entire-leaved form] having been first reported from that state
in 1972 (McGrath & Weedon 1974). In fact, Lactuca saligna is known from
Bryan County, Oklahoma, just north of the Red River border with Texas
(J. Taylor 29740, 28 Aug 1980, BRIT). However, despite this proximity,
the species has not been previously reported from Texas (Correll & Johnston
1970; Stanford 1976; Hatch et al. 1990; Jones et al. 1997).

As part of the collecting effort for the forthcoming Shinners & Mahler s
Illustrated Flora of North Central Texas (Diggs et al.), a collection made in
1998 in Fort Worth (Tarrant County) is apparently the first documented
occurrence of this species for Texas.

Voucher specimen: TEXAS. Tarrant Co.: weedy area in landscape, 301 Crestwood, Fort
Worth, 21 Aug 1998, O’Kennon 14252 (BRIT).

At the collection locality a single large individual was observed as a landscape
weed. This plant was nearly two meters tall and more than a meter wide.
Other populations have not been found in North Central ‘Texas, and it is
unclear whether the species will become more widely established. How-
ever, given that the species is known just north of the Red River and that in
some areas of the Great Plains it is rather frequent (Great Plains Flora As-
sociation 1977; Ted Barkley, pers. comm.), it would not be surprising if

O’KENNON et al., Lactuca saligna in Texas 617

Lactuca saligna were already widespread but unnoticed in the state of Texas.
Lactuca saligna can be recognized by the following description (from original
observations and from Radford et al. 1968; Feraékovd 1976; Barkley 1986;
Stebbins 1993): Taprooted herbaceous annual (or rarely biennial?) 0.3—1(—
2) m tall; latex white; stems erect, branched, usually glabrous or with re-
mote bristles; cauline leaves linear to linear-lanceolate, entire to pinnatified
with 1—2(—3) pairs of narrow, sometimes slightly toothed lobes, with sag-
ittate-clasping base; midrib of leaf white, glabrous or with remote bristles
abaxially; inflorescences spike-like panicles with numerous heads; involu-
cres 10-18 mm high in fruit; corollas yellow (with bluish or purplish on
the abaxial side), drying blue; ligules 4.5-5 mm long; body of achenes 3—
3.6 mm long, ca. | mm wide, with 5—9 nerves on each face; beak of achenes
filiform, usually 1.5—3 times as long as achene body; pappus white, the bristles
ca. 4mm long; 2” = 18; flowering Jul—frost. The accompanying illustra-
tion (Fig. 1) is reprinted from Britton and Brown (1913).

While the Tarrant County individual has mostly pinnatifid lower and
middle leaves and entire upper leaves, leaves in this species can vary greatly.
According to Barkley (1986), “Three phases of L. sa/igna based on leaf
morphology occur throughout its range. The phase with all leaves linear
and entire, referred to by Fernald as f. [forma] rwppiana (Wallr.) G. Beck,
will often occur in the same population with the phase that has all leaves
pinnatifid. The phase with lower cauline leaves pinnatifid and upper cauline
leaves linear and entire may be found with one or both of the other phases.”

The six species of Lactuca occurring in the eastern two-thirds of Texas
(excluding the two species limited to the Trans-Pecos) can be distinguished
using the following key modified from those in Radford et al. (1968), Correll
and Johnston (1970), Barkley (1986), and Stebbins (1993):

1. Beak of achenes (connecting achene body and pappus) stout, 0.5—1(—2)
mm long OR absent; corollas blue or white; body of achenes 4—5 mm
lon L. floridana (L.) Gaertn.
. Beak of achenes thread-like, 2-10 mm long; corollas usually yellow (rarely
cream to pink or pale lavender); body of achenes of various lengths.
2. Beak of achenes equal to or conspicuously longer than body of achenes;

body of achenes ca. 1 mm wide, ca. 1/3 as thick as wide, 5- to 9-

nerved on each face; plants annuals (or rarely biennials) with tap-

root; latex white.

3. Leaves linear to linear-lanceolate, 0.3—5 cm wide, with narrow lobes

OR lobes absent (except for basal lobes); margins of leaves or leaf

lobes entire or remotely prickly-toothed; body of achenes with-

out conspicuous bristles at base of beak; flower 8-15 per head;

abaxial midrib of leaves and |

pa

—y

emotely bristly - L. saligna L.
3. Leaves lanceolate to ovate, oblong-elliptic, or ape. 1—10(—15+ 7

cm wide, with broad lobes; margins of leaves or leaf lobes usually

conspicuously prickly-toothed; body of achenes with conspicu-

ous bristles at base of beak; flowers 14-25 per head; abaxial mid-

rib of leaves and lower stems usually with stiff bristles -..--.--+:+:+0+ L. serriola L.

618 Sipa 18(2)

2. Beak of achenes equal to or shorter than body of achenes; body of
achenes 1.6—3 mm wide, very flat, 1- to 3-nerved on each face; plants
biennials with abundant, tufted, branching roots; latex brownish.
4. Corollas very pale yellow to cream, pink, or pale lavender; lower

leaves usually pinnately lobed; Ba see eaves tia
a clasping base; body of ache —2.5 times as long as wide; in-
frequent in sandy eS in eastern Texa

I a Muhl. var. altos (Torr. & A. Gray) Shinners

4. Corollas usually yellow; a leaves pinnately lobed OR not so;
body of achenes 1.5—2 times as long as wide: widespread in vari-
ous soils in Texas.

5. Body of achenes 4.5—5 mm long, the achenes including beak
7-10 mm long; pappus bristles 7-10 mm long; involucres 13-
22 mm long in fruit; upper leaf paar pinnately lobed, the
margins conspicuously prickly-toothec udoviciana (Nutt.) Riddell
Body of achenes 3.5—4.5 mm long, ie achenes including
4.5-6.5 mm long; pappus bristles 5-7 mm long; Savalas
10-15 mm long 1 in fruit; upper leaf blades usually not lobed,
the margins entire to toothed but usually not prickly-toothed
I

jon

LA

canadensis L.

REFERENCES
BarkLey, I.M. 1986. Lactuca. In: Great Plains Flora Association. Flora of the Great Plains.
Pp. 8 838-1 021. Univ. Press of Kansas, Lawrence. (Cites: Dille, D.P. 1976. A revision of
the genus Lactuca in the Great Plains. M.S. thesis, Kansas State Univ., Manhattan)
BRITTON, N.L. and A. Brown, 1898. An a pe ra of ne ey United States,

al nd . 1913. An lstrated fot of the antes United nace Canada and
~~ the British possessions. The New York Botanical Garden, New York.

CorreLl, D.S, and M.C. ee 1970. Manual of the vascular plants of Texas. Texas
Research Foundation, Renner.

C RONQUIST, A. 1980. Asteraceae. Vascular flora of the southeastern United States 1:1—261

Univ. of North Carolina Press, Chapel Hill.

Diccs, G.M. Jr., B.L. Lirscoms, and R.J. O7KENNON. Shinners & Mahler's illustrated flora
of North Central Texas. To be published in early 1999 by the Botanical Research Insti-
tute of Texas, Fort Worth.

FERAKOVA, V. 1976, Lactuca. In: '1.G. Tutin, V.H. Heywood, N.A. Burges, D.M. Moore,
D.H. Valentine, $.M. Walters, and D.A. Webb, eds. Flora Europaea, Vol 4, Plantaginaceae
to Compositae. Cambridge Univ. Press, Cambridge, England,

FERNALD, M.L. 1950. Gray’s manual of botany, 8th ed. Reprinted eo Dioscorides Press,
Portland, OR.

GLeASON, H.A. 1952. The new Britton and Brown illustrated flora of the eal
United States and adjacent Canada, 3 vols. New York Botanical Garden, Bronx.

and A. Cronquist. 1963. Manual of the vascular plants of ponte United
States and adjacent Canada. Van Nostrand Reinhold Company, New

and . 1991. Manual of the vascular plants of northeastern United States and
adjacent Canada, 2nd ed. Van Nostrand Reinhold Company, New Yor

GREAT PLAINS FLORA AssociaTION. 1977. Atlas of the flora of the a Paine lowa State
Univ. Press, Ames.

a

O’KENNON et al., Lactuca saligna in Texas 619

Haren, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of
Texas. Texas Agric. Exp. Sta. Misc. Publ. No. 1655.

Jones, S.D., J.K. Wiprr, and RM. Montcomery. 1997. Vascular plants of Texas: A com-
prehensive checklist including synonymy, bibliography, and index. Univ. of Texas Press,
Austin.

Macrarn, L.K. and R.R. WEEDON. 1974. New and interesting plants from the central
plains states. Rhodora 76:489-49 1.

Raprorb, A.E., H.E. AHLEs, and C.R. Bett. 1968. Manual of the vascular flora of the
Carolinas. The Univ. of North Carolina Press, Chapel Hill.

Rosinson, B.L. and M.L. FERNALD. 1908. Gray’s new manual of botany, 7th ed.: A hand-
book of the Aas plants and ferns of the central and northeastern United States
and adjacent Canada. American Book Company, New York.

SMALL, J.K. 1903. Flora of the southeastern United States. Published by the author, New

r
: 1913. Flora of the southeastern United States, 2nd ed. Published by the author,
New York.
. 1933. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel
Hill.

SmirH, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2nd ed.
Published by the author, Fayetteville, AR

STANFORD, J.W. 1976. Keys to the vascular plants of the Texas Edwards Plateau and adja-

cent areas. Published by the author, Brownwood, TX

STEBBINS, G.L. 1993. Lactuca. In: ].C.Hickman, ed. ihe oe manual: Higher plants of

California. Pp. 296, 301. Univ. of California Press, Berke

STEYERMARK, J.A. 1963. Flora of Missouri. The Iowa State Univ. Press, Ames.

STRAUSBAUGH, P.D. and E.L. Core. 1978. Flora of West Virginia, 2nd ed. Seneca Books,
Inc., Grantsville, West Virginia.

Taytor, R.J. and C.E.S. Taytor. 1994. An annotated list of the ferns, fern allies, gymno-
ee ms cy flowering plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Univ.,
Dur

WATSON, us. and J.M. Courter. 1889. Gray's manual of the botany of the northern United
States, including the district east of the Mississippi and north of North Carolina and
Tennessee, 6th ed. American Book Co.,

BOOKS RECEIVED

SAMUEL SCHMIDT and GLoria P. Macias, eds. 1997. United States-Mexico
order Environmental Directory. Directorio Ambiental Fronterizo
México-Estados Unidos. 1996. (ISBN 968-6287-94-9, pbk). Universidad
Autonoma de Ciudad Juarez, Lopez Mateos Norte Num. 20, 32310
Ciudad Juarez, Chihuahua, Mexico. Price not given. 389 Pp.

Contenido: Introduccion, Dineci one; Indice a nome en orden alfabético (Gobierno/

Government; ¢ ntal Organizations; Negocios/
Business; Cavecd aeons vem eee Estados/States; hia coaa/iashanions and Miscelane al
Miscellaneous.

S.D. Jones, J.K. Wiper, and RM. Montrocomery. 1997. Vascular plants
of Texas: A comprehensive checklist including synonymy, bibliog-
raphy, and index. (ISBN 0-292-74044-1, hbk). Univ. of Texas Press,
P.O. Box 7819, Austin, TX 78713-7819. $55.00. 404 Pp

Contents: Preface; core ae Introduction; Format; Scientific Names: Species and

Infraspecific Rank; Authors axa; Common Names; Abbreviations and Special Desig-

nations; Conserved Names; C ultivated Plants; Endangered and Threatened Taxa; Federal

xious Weeds; Hybrid ‘Taxa; Misapplied Names; Orthography; State Symbols; Synonyms;

Unsub slished Names; Summary of Taxa; Summary Table; Checklist; Bibliogr, aphy; and Index.

SIDA 18(2): 620. 1998

PENSTEMON METCALFEI
(SCROPHULARIACEAE), A VALID SPECIES

THOMAS K. TODSEN

Department of Biology
New Mexico State University
Las Cruces, NM 88003, USA

In 1905, O.B. Metcalfe collected a Penstemon under spruce at the Look-
out Mine in upper Trujillo Canyon in the Black Range of southern New
Mexico (Metcalfe 1605, NMC!, NY!). Wooton and Standley (1909a) de-
scribed it as Penstemon puberulus Woot. & Stand. Then, realizing that the
specific epithet was previously used, they (1909b) proposed the new name,
Penstemon metcalfei Woot. & Stand. Keck (1945) decided that 2 metcalfei
was synonymous with Penstemon whippleanus Gray, based on his examina-
tion of the NY specimen cited above. His only reservation was that the stem
was puberulous, which differed from the typically glabrous stem of ? whippleanus.

In 1967, I collected a Penstemon under spruce beside the Black Range
Crest Trail on Cross O Mtn ( 7odsen s.n., NMC) and determined it to be P
metcalfei using Wooton and Standley’s “Flora of New Mexico” (1915). This
was the second time the taxon had been collected. Only later did I become
aware that Keck had submerged it under 2 whippleanus.

In the summer of 1995, I collected plants from the Cross O Mtn. Loca-
tion (Todsen 95-7-1, NMC) and compared the live material with the type
and isotype sheets at NMC. There was no significant morphological dif-
ference in plants from the two locations. At the same time, | collected P

whippleanus (lodsen 95-7-7, NMC) from the Sandia Mountains, the type

Taste 1. Morphological differences between P metcalfer and P whippleanus.

Characteristics P. metcalfei P. whippleanus

Stem Puberulous Glabrous

Leaves Entire ‘Toothed

Corolla Pale blue-lavender Deep red-purple
Slightly gibbous Abruptly gibbous

Lower Lip Wider than long Longer than wide

Staminode Not or slightly exserted Well-exserted
Bearded 1/2 its length Glabrous or with tuft

Stamens Not explanate Explanate

Sipa 18(2): 621-622. 1998

622 Sipa 18(2)

locality, and compared that with the Cross O Mtn. Plants. Differences between
the two are shown in Table 1. These characteristics put 2 metcalfei in Keck’s
subsection Graciles rather than in subsection Humiles where he had placed
P whippleanus.

Later, when Crosswhite (1965a) was revising Penstemon section Penste-
mon, approximately equivalent to Keck’s Penstemon section Spermunculus,
he placed P whippleanus in series Whippleani and retained series Graciles,
equivalent to Keck’s subsection Graciles. Crosswhite (1965b) also addressed
a situation in series Graciles, where he divided the series into two “alliances,”
Graciles and Oliganthi, based on differences in distribution, habit, and habitat.
Penstemon metcalfei falls within “alliance” Oliganthi as Crosswhite defined it.

I thank Richard Spellenberg for his many comments and suggestions.

REFERENCES

CrossWHITe, ES. 1965a. Subdivisions of Penstemon section Penstemon. Sida 2:160-162.
965b. Revision of Penstemon section Penstemon Ay emuaaail Il. A western
alliance in series Graciles. Amer. Mid]. Naturalist 74:42°
Keck, D.D. 1945. Studies in Penstemon. VIII. A ono account of the section

ieee Amer. Midl. Naturalist 33:128-2
Wor , E.O. and P.C. StTanb.ey. 1909a. Plants a New Mexico. Bull. Torrefy Club
36:1 12.
and___. 1909b. A new name—VPenstemon metcalfer Wooton & Standley. Torreya
9:145

and . 1915. Flora of New Mexico. Contr. U.S. Natl. Herb. 19:579—586.

COMMENTS ON THE DISTRIBUTION OF
SEDUM PULCHELLUM (CRASSULACEAE)
IN TEXAS

H.L. WHITE, J.R. BRANCH and W.C. HOLMES

Department of Biology
Baylor University
Waco, TX 76798-7388, U.S.A.

J.R. SINGHURST
Wildlife Division
Texas Parks and Wildlife Department
Austin, TX 78744, U.S.A.

Sedum pulchellum Michx. is usually a succulent annual, but may occasion-
ally be a perennial due to the production of secondary roots (Clausen 1975).
Leaves of rosettes, when present, are spirally arranged, spatulate-linear, obtuse
and slightly papillose at the apex. Leaves of floriferous stems are linear and
subterete. Among the North American species, S. pulchellum is unique in
that the leaves of floriferous stems have basal sagittate spurs (Clausen 1975).
Flowers are pale pink, purplish-white, or white, 4-merous, sessile or subsessile,
and occur primarily in 3-branched cymes. Peak flowering takes place in
May and June, but sometimes as early as March.

According to Clausen (1975), Sedum pulchellum occurs on flat rock for-
mations (typically limestone, sandstone, or chert) from NC Kentucky south
to NW Georgia, west through N Alabama to Arkansas, Texas, Oklahoma,
SE Kansas, Missouri, and S Illinois. Northern Mississippi is not included
in Clausen’s distribution, but the species is known from Oktibbeha County
[Brent 181 (IBE!); Stauffer s.n. (MISSA!)]. Recent collections of the species
in the central part of the state and the lack of satisfactory documentation
of its occurrence have prompted us to investigate the distribution in Texas.
Correll and Johnston (1970) state that S. pulchellum occurs “In seepage on
and about calcareous rocks in e. Tex...,” while Hatch et al. (1990) list the
species as a native plant of the Pineywoods and Blackland Prairies vegeta-
tional regions of the state. The species is not included in Shinners’ Manual
of the Flora of North Central Texas (Mahler 1988), although Clausen (1975)
indicated its occurrence within the area treated by the work.

Clausen’s (1975) distribution map gives the most thorough account of
Sedum pulchellum in Texas. Unfortunately, his treatment presents certain

SIDA 18(2): 623-626. 1998

624 Sipa 18(2)

problems. Foremost is the lack of an exsiccata list used in preparation of
the map. He mentions that “A person desiring accurate information about
species in any area not surveyed in detail in the present study should go to
the area himself. The indication on maps that a species occurs in such an
area should be sufficient notice.” The scale of the map is such that it is dif-
ficult to determine the location of the sites. Additionally, points that could
have been included, based upon dates of collections and presence in her-
baria that were cited as being consulted, do not appear. This includes the
Weches formation of San Augustine and Sabine counties in east Texas, an
area from which the species is well known, and Grayson County in NC
Texas. Nonetheless, after canvassing the herbaria cited by Clausen, it was
possible to later verify all but two of the locations indicated on his map
(see Fig. 1). A search of Clausen’s documents in the Cornell University Archives
yielded no additional information. These undocumented locations include
one point in NE ‘Texas, which, as best can be determined, is either in Red
River or Bowie County. Considering the species documentation in nearby
Lamar County and adjacent Oklahoma [McCurtain, Choctaw, and Bryan
Counties (McGregor et al. 1977)], this particular occurrence is of minor
significance. [he other undocumented location given by Clausen, how-
ever, appears to be either in Burnet County or Llano County (or Blanco
County?), an area visited by Clausen in 1970 (Clausen 1975). This uncon-
firmed record is far more notable because it appears to be on the Edwards
Plateau, and therefore may be the most western point in the distribution
of the species

Additional field collections, a survey of herbarium specimens, and Clausen’s
(1975) distribution map in The Sedum of North America were used to document
the expansion of the known distribution of Sedum pulchellum to include
other counties of the Blackland Prairies and Pineywoods as well as the Post
Oak Savannah, Cross Timbers and Prairies, and the Edwards Plateau.

Specimens examined: TEXAS. Bell Co.: Owl Creek Mts., ca. 0.8 air mi SSW of jct. of
Preachers Creek and Owl Creek, Ft. Hood, 4 Jun 1992, Carr 11984 (TEX); N side of
Leon River ca. 0.7 mi ESE of center of Belton Dam, | May 1992, Carr 11868, Enquist &
McNeal (TEX). Bosque Co.: at intersection of Brazos River & Cedar Creek, 97.32808 lat.
31.78761 long., 31 May 1998, Branch 300 (BAYLU). Coryell Co.: Bluff over Leon River
ca. 200 m W of Mother Neff State Park on property of Charlotte & Larry Weiss, 18 May

1996, Holmes 8240 (BAYLU). Grayson Co.: E of Denison in field near creek running

into Smith Creek, 18 May 1950, Gentry | et 1430 (TEX). Lamar Co.: 2.25 mi N of Roxton,
21 May 1937, Cory 23107 (GH). Limestone Co.: Hwy 14 and Pk. Rd. 35, Jun 1993,
Singhurst 1665, Jones, & Blair (BAYLU). ne Pinto Co.: 3. : mi N of jet. Hwy 287 & FM
3027 on Hwy 287, 1.7 mi WSW of Grantham Cemetery, 11 Apr 1998, ad 6593
(BAYLU). Sabine Co.: Weches Formation ca. 4 mi W of ne 9 Apr 1964, Correll &
Correll 29088 (LL); ca. 1 mi W of jet. Hwy 21 and FR 226 on Hwy 21, 31 Mar 1980,
Nixon 10116 (ASTC); 1.4 mi E of jet. of Hwys 21 and 330, 12 May 1980, Nixon 10534
(ASTC); on W side of TX 21 ca. 1.5 mi S of Geneva, 22 Ape 1987, Orzell 5104 & Br idges
(TEX). San Augustine Co.: Weches formation ca. | mi SE of San Augustine, 20 Apr 1962,

Wuite et al., Distribution of Sedum pulchellum 625

me Pes
LPP ROPES ee

Fic. 1 Documented distribution of Sedum pulchellum by county in Texas. The closed circles
represent locations cited by Clausen (1975) that could not be verified (see text for addi-
tional comments).

Correll 25003 (LL); old Hemphill Rd. just E of San Augustine, 10 May 1969, Correll 37202
(LL); ca. 0.5 mi E of San Augustine on Hwy 21, S ona dirt road ca. 1.0 mi, 11 May 1979,
Nixon 9473, Marietta & Matos (ASTC); ca. 1.0 mi S of Chapel Hill on Sanitary Landfill
Rd., 15 Apr 1985, Nixon 14423 (ASTC); 0.2 mi SW of road to Sunrise, 0.9 mi SE of jct.
TX 21 at point 1.1 mi E of center of St. Augustine, 22 Apr 1987, Orzell 5109 & Bridges
(TEX). Travis Co.: near Jester Blvd., Jester Estates, Austin, 1 May 1995, Miller s.n. (TEX);
Bull Creek, 11 Apr 1914, Young s.n. (TEX

We wish to thank the curators of the following herbaria for supplying
the specimens and other information that made this study possible: ASTC,
BH, BM, CHRB, CLEMS, DUKE, E, GA, GH, IBE, K, MISSA, NLU,
NY, NYS, PH, TEX, and US. Assistance provided by Thomas Clausen,
the son of the late R.T. Clausen, and Nancy Dean of the Cornell Univer-
sity Archives was also greatly appreciated.

626 Stipa 18(2)
REFERENCES

Ciausen, R.T. 1975. Sedum of North America north of the Mexican Plateau. Cornell University
Press: Ithaca, New York.

Corrett, D.S. and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. The
ee of Texas at Dallas: Richardson.

Haren, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the idee ls of Texas.
‘Texa one: Exp. Sta. MP-1655, Texas A&M University, College Statio

Mauter, W.E 1988. Shinners’ manu: 7 of the North Central Texas flor a. ae al Research
Institute of Texas, Inc.: Fort

McGrecor, R.L. 1977. Atlas of ie flor a of the Great Plains. lowa University Press: Ames.

AGAVE PETROPHILA GARCIA-MEND. & E.
MARTINEZ, NOM. NOV. (AGAVACEAE)

ABISAI GARCIA-MENDOZA

Jardin Botdnico, U.N.A.M.
A.P 70-614, Del Coyoacdn
04510 México, D.F., MEXICO

ESTEBAN MARTINEZ SALAS

Depto. de Botdnica, eae de Biologia, U.N.A.M.
A.P. 70-233, Del. Coyoacdn
04510 ae D.F., MEXICO

Agave petrophila Garcia-Mend. & E. Martinez, nom. nov. BASONIMO: Agave
gracilis Garcia-Mend. & E. Martinez, Sida 18:227. 1998. Non Agave
gracilis Jacobi, Abh. Schles. Ges. Vaterl. Cult. Abth. Naturwiss. 2:66.
1871. Nombre basado en una cultivada por van Houtte en Gante.
Non Agave gracillima A. Berger, Die Agaven, 33, 288. 1915. BASONIMO:
Agave gracilis A. Berger, Die Agaven, 33. 1915. = Manfreda elongata
Rose.

Agave gracilis Garcia-Mend. & E. Martinez es un homénimo posterior a
las propuestas del mismo nombre por Jacobi, 1871 y Berger, 1915, por lo
que con base en el articulo 58.1 del International Code of Botanical No-
menclature (1994), se reemplaza este nombre con un nuevo epiteto.

REFERENCIA

GarciA-M., A. y E. MARTINEZ. 1998. Una nueva especie de Agave, subgénero Littaea (Agavaceae)
de Guerrero y Oaxaca, México. Sida 18:227—230.

SIDA 13(2)5 627 01998

AMELIA ANDERSON LUNDELL (1908-1998)
BOTANICAL ARTIST AND FRIEND

RUTH ANDERSSON MAY
4920 Mangold Circle
Dallas, TX 75229, U.S.A.

Amelia Anderson Lundell, born December 21, 1908 in Fort Worth, who
passed away in Dallas in her 90th year on August 27, 1998, gave much to
the world of art and science. The Botanical Research Institute of Texas (BRIT)
has many of her renditions of shells and wildflowers that she donated after
the death of her husband, Cyrus Longworth Lundell, the posthumous re-
cipient of BRIT’s International Award of Excellence in 1994. She drew all
the insects pictured in the Dallas Planting Manual. Amelia’s Sand-Verbena
(Abronia ameliae Lundell, Verbenaceae) is named for her. She and Cyrus
found the undescribed endemic Abronia near the Texas coast on a collect-
ing trip years ago. Many of her paintings of wildflowers are in a suite of
rooms near the Rare Book Library of UT Austin. Amelia made exquisite
needlework embroideries and enjoyed sewing for her granddaughters when
they were babies. Her home was adorned with her artwork and she loved
to cook. The following is one of her favorite recipes, always included in
her collection of home baked Christmas cookies that were shared with friends.

AMELIA’S BUTTER FINGERS
2 cups flour
lL cup butter
1/2 cup whipping cream (not whipped)
Blend flour and butter. Add cream. Roll out and fold over, dusting lightiy
with sugar, four or more times. Cut in sticks. Brush top with egg white
and sprinkle with sugar. Bake at 300 degrees for at least 20 minutes or more.

She was an honorary member of Founder’s Garden Club (GCA) of Dallas, the
Dallas Woman's Club, and the Dallas Garden Club. Her generosity made pos-
sible the handsome gift of the Wi/dflowers of the United States (Rickett 1966-
1973, vols. 1-6) series of books to the Woman’s Club library. This is a lasting
memorial to a beloved, knowledgeable, and talented friend of botany.

Obituary
LUNDELL, AMELIA ANDERSON. Dallas Morning News. Obituaries, 34 A. Saturday,
29 August 1998.

Sipa 18(2): 629-630. 1998

630 Sida 18(2

SS

‘00a
“Steen ean,

ee

ia Anderson Lundell at BRIT’s International Award of Excellence in 1994 where her

late husband was honored.

Ame

Ardisia ellipticifolia Pipoly & Ricketson, nom. nov. 512

Ardisia eucuneata (Lundell) Pipoly & Ricketson, comb, nov. 512

Ardisia feniana Pipoly & Ricketson, nom. nov. 512

Ardisia guanacastensis (Lundell) Pipoly & ah comb. nov. =

Ardisia guinealensis (Lundell) Pipoly & Ricket

Ardisia hatoana (Lundell) Pipoly & Ricketson, oan nov, 515

Ardisia heterotricha (Lundell) Pipoly & Ricketson, comb. nov. 312
Ardisia hornitoana Pipoly & Ricketson, nom. nov. 513

Ardisia hugonensis (Lundell) Pipoly & Ricketson, comb, nov, 513

ketson, comb. nov. 313

on, comb, no

Ardisia ixcanensis (Lundell) Pipoly & Ric

Ardisia jaliscensis (Lundell) Pipoly & Ricketson, comb. nov. 513
Ardisia knappii (Lundell) Pipoly & Ricketson, comb. nov. 413
Ardisia lajana cael Pipoly & Ricketson, comb, nov. 515

Ardisia latisepala (Lundell) Pipoly & Ricketson, comb. nov. 513
Ardisia leptopoda (Lundell) Pipoly & Ricketson, comb. nov. 513
Ardisia mesoamericana | a & Ricketson, nom, nov. 315

Ardisia m rdeana (Lundell) Pipoly & ne comb. nov. 513
Ardisia ne a
Ardisia neomirandae Pipoly & Ricketson, nom. nov. 514
Ardisia niambiensis Pipoly & Cogollo, sp. nov. 390
Ardisia oaxacana (Lundell) Pipoly & Ricketson, comb. nov. 314
oe opegrapha Ocerst. subsp. paquitensis (Lundell) Pipoly & Ricketson, comb et stat. nov, 457
pha Ocrst. subsp, wagneri (Mcz) Pipoly & Ricketson, comb. et stat. nov. 452

losa ( aie Pipoly & Ricketson, comb. nov

_

Ardisia patvidentculata Pip ly & Ricketson, nom. nov.
Ardisia parvipunctata (Lundell) Pipoly & Ricketson, cone nov. 514
Ardisia samalana (Lundell) Pipoly & Ricketson, n, comb. nov, 514
Ardisia steinii Pipoly & Ricketson, nom. nov, 514

Ardisia stevensii (Lundell) Pipoly & Ricketson, comb. nov. 314
Ardisia toroana (Lundell) Pipoly & Ricketson, comb. nov. 514
Ardisia triangula (Lundell) Pipoly & Ricketson, comb. nov. 514
Ardisia utleyi peer pe & Ricketson, ee nov. 515

Ardisia warneri Pipoly & Ricketson, nom.
Ardisia wendtii (undell) Pipoly & Ricketsor seit nov. 515
x78

Arnoglossum album L.C. Anderson, sp. nov. 378
Berlandiera res (Mics: ) Nutt, var. scabrella Nesom & Turner, var. nov. 499
OS

Clusia aristeguietae (Maguire) Pipoly, comb. 1

Clusia Senay Pipoly & Cogollo, sp. nov a
sein comans (Mart.) Pipoly, comb, nov. 408

ta (Cuatrec.) Pipoly, comb. nov. 408
Clusia ene a (Cuatrec.) Pipoly, comb. nov. 408
Clusia mirandensis (Cuatrec,) Pipoly, comb. nov. 405

Clusia niambiensis, Pipoly, Cogollo et Gonzalez, sp. nov. 396
104

520

ee isi saes Pi tpoly & Cogollo, sp. nov
C Hameed & Madhu isoodanan sp. nov,

Malaxis palustris Espejo & Lopez-Ferrari, sp. nov, 412
Malaxis alvaroi Garcia-Cruz, R. Jimenez & L. Sanchez, sp. nov. 414
Myrsine calcarata (Lundell) Ricketson & Pipoly, comb. nov. 506
Myrsine juergensenii (Mcz) Ricketson & Pipoly, comb. nov. 506

er an eunne ense Nesom, sp. nov. 523
ckera bolanderi var. harfordii (Greenm.) D.K. Trock & TM. i Day comb.nov, 386
Packera sinh ener evs M. . lev) D.K. Trock a TOM. Barkley, comb. nov. 386
Packera dimorphophylla var. paysoni (1M. Barkley) D.K. Tr Sie & TM. Barkley. comb. nov

ee nov. 35
M. Barkley, comb. nov. 386

Packera pse ay ~ flav ne K. Trock & TM. Bar

Packera pseudaurea var. semicordata (Mack & Bush) D.K. Trock &
metcalfei (Greene) D.K. Trock & TM. Bar ee comb, nov. 2!

Packera neomexicana \
Barkley, comb. nov. 386

Packera neomexicana \: ee (Greene) D.K. Trock & TM.
Packera i D.K. Troc f & TM, Barkley, nom. nov. 387
Soimaihes iceloe Solano & Garcia-Mend., sp. nov.

LSS)

Agalinis (Scrophulariaceae) in the Ozark Highlands
Jobn F. Hays

Flora ornamental en plazas y jardines publicos del area metropolitana de Monterrey, México
Alejandra Rocha Estrada, Teresa FE. Torres Cepeda, .

Va. del Consuelo Gonzalez de la Rosa, Salomon J. Martinez Lozano y
Marco A. Alvarado Vazquez
579

Documented chromosome numbers 1998:1, Chromosome numbers in Carex section Ovales (Cyperaceac)
Variations, and corrections

PLE. Rothrock and A.A. Reznicek

sus

- Additions,

Vascular plants new to Kans:
ral Mg C. Freeman and see VWeGregor

New and noteworthy angiosperm records for lowa
Craig C. Freeman

605

lziola | i P > Oryzeae) previously unreported from Texas and a key to Texas species
Gephan Hatch, David J. en dnd James A. Thomas
oll

Lactuca saligna (Asteraceae), @ lettuce new for Texas
Robert J. O'Kennon, George M. Diggs, Jr. and Barney L. Lipscomb
615

Penstemon metcalfei (Scrophulariaceae), a valid species
Thomas kK. Todsen
621

Comments on the distribution of Sedum pulchellum (Crassulaceae) in Texas
FLL. White, [.R. Branch, W.C. Holmes, and JR. Singburst
623

Agave petrophila Garcia-Mend. & EF, Martinez. nom, nov,

(Agavaceae)
Abisat Garcia-Mendoza y Esteban Martinez Salas
627

In Memoriam—Amelia Anderson Lundell (1908-1998)
Ruth Andersson May
029

Book notices and reviews 388, 394, 400, 410, 432, 478, 492, 518, 578, 610, 620

Index to new names and new combinations in this issue

Agave petrophila Garcia-Mend. & FE. Martinez, nom. nov. 027

Ardisia alajuelae (Lundell) Pipoly & Ricketson, comb. nov. 511

Ardisia albipedicellata Pipoly & Rickctson, nom. nov. 511

Ardisia albipetala (Lundell Pipoly & Ricketson, comb. nov. 515

Ardisia albisepala (Lundell) Pipoly & Ricketson, comb. nov, 51]

Ardisia azaharensis Pipoly & Ricketson, nom. nov. 511

Ardisia bartletii Lundell subsp. lilacina (Lundell) Pipoly & Ricketson, comb. ct sta
ae a ne & Ricketson, nom. nov. 511

nov, 4

undell) Pipoly & Ricketson, comb. nov. 511
Ardisia dressledi & Ricketson, nom. nov. 511

sia dryeri Pipoly & Ricketson, nom. nov. 512
Ardisia duripetala (Lundell) i ly & Ricketson, comb. nov. 515
Ardisia eciliata (Lundell) Pipoly & Ricketson, comb. nov. 512

1O

(continued on tside back cover) ISSN OQU36-1488

CONTENTS

Villasenoria (Asteraceae: Senecioneae): A new genus and combination from Mexico

Bonnie L. Clark

31

-ackera subnuda comb. nov., a corrected name for Packera buekii (Asteraceae: Senecioneae)
Debra K. Trock and Theodore M. Barkley

35

\ new species of Phaseolus (Fabaceae) from West- oes Mexico
RLY’ mundo Ramirez-D lg wdillo and Alfonso Del, gd li

37

nas
Salt TLS

bs oe of the American black-fruited hawthorns Crat da, C, rivularis,
». Saligna and C. brachyacantha to C. ser, Dolighisianae (Rosacese

B. Phipps:
47

th
7

4 /

he segregates of Senecio, s.|., and Cacalia, s.|., in the flora of North America north of Mexico
heodore M. Barkley
6

_ new variety of Physaria newberryi (Brassicaceae) from New Mexico
‘obert C. Sivinski

73
ynopsis of New World Commicarpus (Nyctaginaceae

eil A. Harriman

79

fuhlenbergia jaime-hintonii (Poaceae: Chloridoideae), a new species from Nuevo Leon, Mexico
aul M. Peterson ee V Sei

85

new species of Emorya.(Buddlejaceae)

fark H. Mayfield

93

iscovery Of the Indo-Malesian genus Hymenandra (Myrsinaceac) in the Neotropics, nd its
oreotropical implications

hn J. Pipoly IT and Jon M. Ricketson

1

lyrsine luae (Myrsinaceae), a new species from Brazi
in M. Ricketson an 1. je hn J. Pipoly It
(7

otanical results from the 1995 Bismarck-Ramu Expedition in Papua New Guinea
’ Takeuchi

5

udies in New World Amaranthus (Amaranthaceae)
imes Henrickson

wo new species of Tripogon (Poaceae) from India CONTRIBUTIONS
K. Pradeep and C.N. Sunil TO BOTANY
)9

arnebydendron, a new generic name (Fabaceae, Caesalpinioideac| Detatiead, Brownea group)
seph H. Kirkbride, Jr.
15
VOLUME 18
ontinued on back cover) NUMBER 3
SEPTEMBER, 1999

CONTRIBUTIONS TO BOTANY
FOUNDED |
LLOYD H. anes
1962

Wm. 7 Mahler
Publisher 197 -199?2

Barney L. Lipscomb
Editor
Botanical Research Institute of Texas, Inc.
509 Pecan |
Fort Worth, Texas 7610 ), USA
817 332-4441 / 817 a4 1] : FAX
Electronic mail: sida@brit.org
Home page at the URL: http://www.britorg/sida

John W. Thieret Prof. Dr. Felix Llamas
Associate Editor Contributing Spanish Editor
Biological Sciences Dept. Dpto. de Botanica, Facultad de Biologia
Northern Kentucky University Universidad de Leon
Highland Heights, Kentucky 41076, USA -24071 Leon, Spain

Guidelines for contributors are available upon request

nd on the inside back cover of the last issue of each volum
pete per year: $25. Individual, $45. USA Institutions, $50. ont USA;
num

ders issued twice a year

a)
an

© SIDA, CONTRIBUTIONS ae ANY, Volume 18, Number 3, pages 631-940
ypyright 1999
Botanical Rese: ee) Insutute of Texas, Inc.
Printed in the United States of Americ
ISSN 0036-1485

VILLASENORIA (ASTERACEAE: SENECIONEAE): A
NEW GENUS AND COMBINATION FROM MEXICO

BONNIE L. CLARK'!
Herbarium, Division of Biology
Kansas State University
Manhattan, KS 66506-4901, U.S.A.

ABSTRACT
A new genus, Villasenoria (Asteraceae: Senecioneae), is described and a new combina-
tion, Villasenoria orcuttii (Greenm.) B.L. Clark, comb. nov., is proposed.

RESUMEN

Se describe un nuevo género, Vil/lasenoria (Asteraceae: Senecioneae), y se propone una
nueva combinacion, Véllasenoria orcuttii (Greenm.) B.L. Clark, comb. nov

During the course of revising the traditional section Terminales Greenm.
of the genus Senecio (Clark 1996), I had the opportunity to see an under-
collected entity originally described by Greenman (1912) as Senecio orcuttii.
This entity was subsequently included in the segregate genus Telanthophora
by Robinson & Brettell (1974). With the collection of more specimens and
better habitat information, it became evident that the entity is sharply dis-
tinct from the rest of Telanthophora and other groups of Senecio, s.1., and that
it should be placed in its own genus. Vi//asenoria is described herein as a
new genus to accommodate Senecio (Telanthophora) orcuttit.

The plants of the traditional section Terminales are shrubs and small trees
mostly of middle elevations and the highlands of Mexico and Central America.
As conceived by Greenman (1901), this section is distinguished by plants
with stems that are abruptly contracted or foreshortened just below the terminal
capitulescences, which are pedunculate clusters of corymbiform cymes. Studies

obinson and Brettell in the 1970s led them to suggest that the Mexi-
can and Central American species of the super-genus Senecio should be split
into several segregate genera, and Senecio sect. Terminales Greenm. was split
into two genera, Pzttocanlon and Telanthophora (Robinson & Brettell 1973,
1974). Though slow to receive support in the literature, recent synthetic
publications, e.g., Jeffrey (1992), Bremer (1994), and Barkley et al. (1996),
have adopted these segregate genera.

Senecio orcuttit is an anomaly in either Te/anthophora or Pittocaulon in that
it has large principal leaves (to 7 dm long) that are pinnately compound,
and the stems merely taper to the capitulescence. Moreover, the capitulescence

'Present Address: 8201 Hauser Drive, Lenexa, KS 66212-2542, U.S.A.

Sipa 18(3): 631-634. 1999

632 Sipa L8G)

is a loose, elongate cluster of numerous heads, without the distinctive corymbiform

or dome-shaped aspect of the capitulescences of Te/anthophora and Pittocauton.
The species is known only from limestone karst outcrops in rainforests from
LOO to nearly 2000 m elevation in Chiapas, Oaxaca, and Veracruz, Mexico.
On the other hand, species of P/trocai/on occur in seasonally dry scrublands
of central and southern Mexico at 300-3250 m, while species of Telantho-
phora occur in cloud forests or ae hardwood forests from central Mexico
southward to Honduras, at GO0—SO000 m.

Pittocaulon, Telanthophora, me the new genus Vi/lasenoria are separated
by the following key

KEY TO GENERA
1. Leaves pinnately compound, the largest to 7 dm long overall; capitulescences
elongate and neither corymbiform nor dome-shaped; plants single-stemmed,
the stems tapering to the capitulescences; plants mei achene surfaces
papillace. Villasenoria

|. Leaves variously entire to lobed bue not compound, rarely more than 4 dm
long overall; capitulescences of terminal, dome-shaped corymbiform cymes;
plants single stemmed or multi-branched, with stems abruptly foreshort-

f: -es smooth,

ened below the capitulescence; shrubs or small trees; achene s
glabrous.
2. Leaves palmately veined, seasonally deciduous, mostly absent at Howering
time; stems rubbery and subsucculent, with subepidermal resin ducts; pith
chambered. Pittocaulon

2. Leaves pinnately veined, persistent; stems woody, without evident sub

epidermal resin ducts; pith continuous. Telanthophora
Villasenoria B.L. Clark, gen. nov. Type species: Vi/lasenoria orcuttit (Greenm.) B.L. Clark.

Terbae lignescentes, caule e radicibus fascicul fibrosi i Apicem versus in inflorescentiam

terminalem, corymbiformem vel paniculatum abeunti. Foloia pinnatim ae in-
fra inflorescentiam Seimei C apie numerosa radia ata; pay laria (plus/minus) 8,
li iis subtribut Tussalagineorum

aequilonga, bisertata; f
tribui Senecionearum congrui, superficie stigmatica trans interiorem styli ramulorum faciem

I

continua; antherarum collum parum incrassatum sed cellulae basi haud auctae.

Plants erect, to 3.5 m tall, stems single, arising from a cluster of fibrous
roots; leaves present just below the capitulescence, lower leaves deciduous.
Stems terete with solid pith, glabrous but with circular to horseshoe-shaped
leaf scars; bases of recently dropped leaves persistent; stems tapering to the

capitulescence and not conspicuously foreshortened. Leaves alternate, pin-
nately compound, petiolate; blades oblong to ovate in outline, leaflets ovate
or obovate to broadly lanceolate; adaxial surface lightly stromose-papillate
to lightly pubescent. Capitulescences terminal, open-paniculiform to loosely

corymbiform clusters of capitula, peduncles multibranching, with scattered
linear-subulate bracts. Capitula numerous, radiate. Involucre narrowly cam-
panulate to cylindrical; receptacle flat, naked or but shallowly alveolate, fisculose;

Ciark, Villasenoria, a new genus from Mexico 633

phyllaries 8(—9), biseriate, in an inner and an outer series, linear-lanceolate,
slightly swollen or thickened at the base, inner phyllaries with scarious margins;
calyculate bracts few and inconspicuous, linear. Ray florets 2—5, ligulate,
pistillate, corollas yellow, tube cylindrical, lamina apex with 3 minute teeth,
veins unbranched. Disk florets 16-23, corollas yellow, gradually to some-
times abruptly expanded upward, lobes 5. Anther bases sagittate, anther
collars slightly swollen but the cells not basally dilated; endothecial cell
walls with radial thickenings. Styles with stigmatic areas entire or weakly
transitional, with morphologically distinctive cells in triangles at the bases
of the stigmatic areas; apices unappendaged, truncate to conical, with fringing
papillae of uniform length; stylopodia free. Achenes cylindrical, glabrous
but with papillate projections; epidermal cells of pericarp oblong in sur-
face view; carpopodia of 5-17 rows of thickened quadrate cells, margins
regular to irregular; ovary walls with heterohexagonal and occasionally with
acerose crystals. Pappus of numerous white, uniform, capillary hairs. Chro-
mosome number unknown.

Etymology.—The genus name honors Dr. José Luis Villasefior, a dedicated
botanist of the Instituto de Biologia, Universidad Nacional Auté6noma de
México, in Mexico City.
bee orcuttii (Greenm.) B.L. Clark, comb. nov. Senecio orcuttii Greenm.

Field Mus. Bot. 2:350. ae di oer orcuttit aia ) H. Rob. & Brectell,
ee 27:428. 1974. Type: MEX . VERACRUZ: Omealca, near Cérdoba, on
imestone cliffs, 6 Apr 191( . a Orcutt 3150 ener eae pe: F!; isorypr: MO!).

Additional specimens examined: MEXICO: Chiapas: de Tuxtla G. a Montecristo (N
Tuxtla G.), 17 Nov 1949, F. Miranda 5695 (MEXU). Oaxaca: San Felipe Usila, en cerro
Casa de San Felipe de Usila, tipo veg. Selva alca perennifolia, suelo negro rocoso, elev. 250

, 17 Oct 1989, J.L. Calzada 14977 (KSC); 5 km N of Huatla de Jiminez on road to Santa
ee Chilchotla, on steep slopes of calcareous rock, in rainforest remains along edge of
coftee parce 18°09'N, 96°52'W, elev. 1200 m, 25 Mar 1992, B.L. Clark, A. Salinas
G J. lL. Calzada 53 (KSC, MEXU); 5 km N of Huatla de Jiménez on road to Santa Maria
Chilchotla, on ae slopes of sn rock, in rainforest remains along edge of coffee
pues 18°09'N, 96°52'w, elev. 1200 m, 16 Mar 1994, B.L. Clark & A. Campos V.

KSG, MEXU), 749 SC, oo 151 (KSC, MEXU); along road from Teotitlan
a Camino to Chilchot'a, 4.4 mi beyond turn-off to H uatla de Jiménez, steep slopes with
huge boulders, a few native trees persisting in a coffee plantation, elev. lane im, 23: Feb
1979, TB. Croat 48378 (KSC, MO): Chilchorla, aa de Jiménez, bosque de encinos,
suelo pedregoso, pardo-obscuro, 1200 msnm, 6 Apr 1975, R. Herndnder 3105 (MEXU
eracruz: Axocuapdn, selva media subperennifolia, elev. 450 m, 30 Dec 1988, ML. Chazaro,
L. Robles, 6 J.L. Tapia 5793 (WIS); Jess Carranza, km 6 del Camino Cedillo-Rio ee.
selva alta perennifolia, cease y secundaria, suelo café claro arcilloso rocoso, 17°10'N,
94°40'W, elev. 150 m, 18 Jan 1975, B. ae 3905 (MEXU): Hidalgocielén, km 3 del
Camino Cedillo-La Laguna, selva alta perennifolia, primaria, suelo c :
elev. 150 m, 19 Jan 1 975, B. Dorantes 3971 (MEXU} Atoyac, 13 — LOS ieee. Mee
1394 (MEXU, MICH); Hidalgotitlan, 3 km SW of Campamento La Laguna, selva alta
perennifolia, original forest on isolated small fills of extreme limestone karst, surrounded

Ww

=

634 Sipa 18(3)

by flat pastures with scattered remnant trees, 17°16'N, 94°32'W, elev. LOO m, 6 Mar 1984,
M. Nee 30011 (KSC, MO, NY); Motzorongo, mountain side, 11 Feb 1892, ).G. Smith 135
(MQ); Hidalgoticlan, 300 m W Campamento Pian: Cedillo, parte alta cantiles Rio
Solosuchil, selva alta eee primaria, suelo carstico, sobre suelo rocoso por completo,

17° 16'N, 94°36'W, elev. 150 m, 1 Nov 1975, B. Vdsques 1632 (MEXU); Coetzala, por el
camino a Ceotzapotitla, sue alta subperennifolia, secundaria, suelo rojo amarillento con
rocas aflorantes, 25 Apr 1976, V. Vedsquez 400 (MEXU); Cérdoba, San Rafael, bosque de
encino en ladera de cerro, Be 900 m, 28 Mar ae : Ventura aria oe auacieern
0.6 km al E del Poblado 7, sobre camino a La Raya, a C rturbada

de Spondias, Bursera, Dai etc., rodeada por iy 17°19'N, 94°31'20" W. ae 130
m, 31 Mar 1981, 7) Wendt, et al. 3122 (MEXU); Minanclan, 13.7 km al E de La Laguna,
sobre la terraceria a Uxpanapa, luego 6.2 km al N sobre el camino nuevo (no completo) a
Belisario me ce selva mediana con Dralimm, en area karstica con afloramientos de
piedra, 17°19'30"N, 94°23'W, elev. 130 m, 11 Feb 1981, T. Wendt, A. Villalobos, & D.
Olmstead 2838 (MEXU).

ACKNOWLEDGMENTS

This study is a portion of a doctoral dissertation prepared at Kansas State
University under the supervision of T.M. Barkley. I wish to thank the cura-
tors of the following herbaria that lent specimens to herb. KSC for this study:
ASU, CAS, DAV, DS, F GH, KANU, LL, MEXU, MICH, MO, MSU, NMC,
NY, UC, UMO, US, and WIS. Studies of collections at BM, K, MEXU and
US helped form a broader picture of the assemblage. Iam indebted to Dr.
José Luis Villaseftor and the staff of the Herbario Nacional (MEXU) at Universidad
Nacional Aut6noma de México for their kind advice and for their assistance
with collecting trips in México. Thanks also to Rupert Barneby of the New

York Botanical Garden for preparing the Latin diagnosis.

REFERENCES

Barkey, T.M., B.L. Clark, and A.M. Funston. 1996, The segregate genera of Senecio, s.L.,
nd Cacalia, s.1., in Mexico and Central America (Asteraceae: Senecioneae). In: D.J.N

an
Hind and H.J. Bee ntje, eds. Compositae: mys tenance’. Proceedings of the Gea cal
pias Conference, Kew, 1994. 1:613—620. Royal Botanic Gardens, Kew, U

Bremer, K. 1994. Asteraceae: eae Timber & Pat ae Oregon.
Crark, B.L. 1996. A eat) of variation in Senecio Sect. Terminates ( Greenm. (C ompositae:
Senecioneae). Ph.D. dissertation, Kansas State University, Manhattan, Kans

GREENMAN, ].M. ae Monssiaptie der nord- und centralamerikanischen Arten = Gattung
Senecio. I. Teil. Allgemeines und Morphologie. Leipzig. 37 pp. Reprinted in Syst. Bot
come 32:1—33. 1902

4 1912 Tyiasanees of new species and notes on other Spas en chiefly
from “Me X1CO ad Central America. Field Mus. Nat. Hist., Bot. Ser. 2:329—350.

Jerrrey, C. 1992. The tribe Senecioneae (Compositae) in the Mascarene Islands with an
annotated worl Id checklist of the genera of the tribe. Notes on Compositae. VI. Kew

ull. 47:49-LO9

ROBINSON, H. and R.D. Brerreti. 1973. yal on the Senecioneae (Asteraceae). I.A new

genus, Prttocaulon. Phytologia 26:45 1 2.
974. Studies in the Soe cae (Asteraceae). co a genera Psacaliopsis,
Banandin: Telansbonbons aad Roldana. Phytologia 27:402—

PACKERA SUBNUDA COMB. NOV.,
A CORRECTED NAME FOR PACKERA BUEKII
(ASTERACEAE: SENECIONEAE)

DEBRA K. TROCK AND THEODORE M. BARKLEY

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
dtrock@brit.org; tharkley@ brit.org

ABSTRACT

The new combination, Packera subnuda (DC.) D.K. Trock & T.M. Barkley is proposed,
based upon Senecio subnudus DC.

RESUMEN

Se propone la nueva combinacién Packera subnuda (DC.) D.K. Trock & T.M. Barkley, en
base a Senecio subnudus DC.

The North American entity once known as Senecio subnudus DC. and later
as Senecio cymbalarioides H. Buek has a tangled nomenclatural history, espe-
cially when transferred to the segregate genus Packera. We reviewed these
complexities (Trock & Barkley 1998) and proposed the new name, Packera
buekit D.K. Trock & T.M. Barkley for this plant. Drs. K.N. Gandhi and John
Strother have both pointed out that the nomenclatural session of the recent
International Botanical Congress (Saint Louis, 1999) passed a rule which
pertains to our nomen novum. The new rule is an addition to the present art.
53.6, & ex. 17, to be included in the forthcoming International Code of
Botanical Nomenclature ICBN). According to the concept of the new rule,
a renamed legitimate homonym has priority over its zomen novum during a
transfer to another genus. In this case, the basionym Senecio subnudus is le-
gitimate in the segregate genus Packera and, therefore, the replacement name,
Packera buekii is superfluous. The following new combination is proposed:
Packera subnuda (DC.) D.K. Trock & T.M. Barkley, comb. nov. Basiony:

Senecio subnudus DC., Prodr. 6:428. 1837 (non Senecio subnudus DC, Prodr. 6:405. 1837,

African plant.). Senecio cymbalarioides H. Buek, Gen. Sp. Synon. Cand. 2:6. 1840
(non pairs cymbalarioides Nutt., Trans. Amer. Phil. Soc. II. 7:412. 1841.). Packera
buekii D.K. Trock & TM. Bele Sida 18:387. 1998 (nom. superfl.).

REFERENCE

Trock, D.K. and TM. Bark.ey. 1998. Seven new nomenclatural combinations and a new
name in Packera (Asteraceae: Senecioneae). Sida 18:385—387

Sipa 18(3): 635. 1999

WS

Sipa 183

BOOK NOTICE

Douctas B. Evans. 1998. The Cactuses of Big Bend National Park. (ISBN
()-292-72098-x, hbk, 0-292-72099-8, eg ) University of Texas Press,
Box 7819, Austin, TX 78713-7819. $12.95. 82 Pp.

This book provides color photographs for che common cacti of this important nacional
park. After a brief introduction and glossary, the book is divided into 12 chapters, arrangec
by genus, including Ariocarpus, Coryphantha, Echinocactus, Echinocerens, Echinomastus, Epithelantha,
Ferocactus, Glandulicactus, Mammillaria, Neolloydia, Opuntia, and Thelocactus. Finally, the book
ends with a list of cacti historically reported but not recently seen, or cacti that one would
sect but have not as yet been found in the park.

The photographs are excellent, and the high quality, glossy paper accentuates tl
thetic value. The descriptions are pointed at the hobbyist but are practical and useful to
everyone. Both common and scientific names are given for each species, and no precise lo-
ities are given, presumably to inhibit poachers. For some species, details about the eco-

eX]
1eir aes-

—

cali

logical life history and elevational range is also giv

is is a wonderful, thrifty “coffee table” and f ‘eld a cuit, and | highly recommend it to
f Texas.—John J.

am

slant enthusiasts planning to visit che park, or working with the flora 0

Pipoly HL.

Sipa 18(3): 636. 1999

A NEW SPECIES OF PHASEOLUS (FABACEAE)
FROM WEST-CENTRAL MEXICO

RAYMUNDO RAMIREZ-DELGADILLO
Instituto de Botdnica, CUCBA
Universidad de Guadalajara
Apartado Postal 139, C. P. 45110
Zapopan, Jalisco, MEXICO

ALFONSO DELGADO-SALINAS

Departamento de Botdnica
Instituto de Biologfa, UNAM
Apartado Postal 70-367, C.P. 04510
México D.F. MEXICO

ABSTRACT
We report the existence of a new species, Phaseolus albescens, from western Mexico, with
close affinities to P. vulgaris, P. coccineus, P. costaricensis, and particularly to P. polyanthus, and
describe plants commonly occurring in wild habitats, examine morphological and anatomical

characters and give notes on observations made on its floral biology. An illustration of the

planc, photographs of its pollen, and stem anatomy are provided. Also, a key to the species
of the Phaseolus vilgaris group is presented.
RESUMEN

Se reporta la existencia de una nueva especie, Phaseolus albescens, del occidente de México,
specs atinia aP. vul, gars, P. coccineus, P. uiatdane y en particular a P. pee yanthus describen
at ae is

L
7 se presentan notas sobre iones efectuadas a su biologia aneae ia especie se ilustra

y se provee de eee a su nae y anatomia del tallo. También, una clave de las especies
del grupo Phaseolus vulgaris es presentada.

INTRODUCTION

Populations of Phaseolus albescens have been recognized as morphologi-
cally distinct for a decade (Delgado-Salinas 1988; Ramirez-Delgadillo 1991).
Some of the differences were clearly detailed by McVaugh (1987), who treated
this species as Phaseolus, aff. P. coccinens for the Flora Novo-Galiciana. In-
deed, McVaugh (1987) recognized Phaseolus albescens as different, but not
convincingly so from P. coccineus. According to him, this plant is easily dis-
tinguished from sympatric wild plants of P. coccinews by the color of its flowers
[“pale-lavender,” “old-rose,” or “pale rose-pink, fading yellowish”}, and by
“... less tendency for the flowering axes to be crowded towards the tip.” He
also compared it with P. polyanthus Greenman (also recognized as P. coccineus

Sipa 18(3): 637-646. 1999

Sipa 18(3)

subsp. darwinianus or also as P. coccinens subsp. polyanthus), stating that this
Nueva Galicia plant has shorter, ovate to lanceolate bracteoles and glabrate
foliage.

During the course of a phylogenetic study of the genus Phaseolus, inferred
from molecular data and non-molecular data (Delgado-Salinas et al. unpublished),
three collections of P. a/bescens from west-central Mexico were shown to be
clearly different from three samples of P. polyanthus, one wild collection from
Guatemala, one cultivated from Puebla, and one escaped from Pert. Phaseolus
albescens collections differ from those of P. polyanthus, by at least 51 pair-
base mutations, 7 of which are transversions, and one deletion of 3 base
pairs. In addition to these molecular characters, morphological differences
observed during previous studies were shown to be distinguishing. Such
characters included the floral bracteoles, flower color, and glabrate foliage.
Also the P. a/bescens clade was restricted to a region north of the range of P.
polyanthus.

In this paper information on the anatomy of the stem and on the floral
biology of this species is brought into consideration and thus, the species ts
described and finally formalized below:
oT ae McVaugh ex R. Ramirez & A. Delgado, sp. nov. (Fig.

. Phaseolus, aff. P. coccinens sensu McVaugh, in McVaugh, Flora ae Galiciana
ee i Type: MEXICO. Jatisco: Municipio de Ciudad Guzman, Naranja
Verde-La Retama, a 2 km sobre la deviacién que va al Fresnito de la carretera Cd.
Guzmian-El Grullo, N 19° 36'59",O 103° 30! 54.7", 1640 m, 19 Nov 1996, Delgado
et al. 1705 (HOLOTYPE: MEXU; Isotype: IBUG)

Phaseolus polyantho Greenman sylvestri affinis, sed differt bracteolis Se brevioribus,
floribus lilacinis, postea albescentibus, demum lutescentibus, ovulis 4—5, et distributione
8COg raphica 1 in Mexico occidentali

Perennial herbaceous and seat vine, with secondary growth developed
in basal stems, 2—3 cm in diameter, with corky appearance and displaying
conspicuous rows of lenticels; root long and lignescent, non-tuberous; stems
up to 10 m long, terete, leaning or twining, sparingly branched, covered
with sparsely appressed and uncinate hairs. Leaves membranous and large,
up to 30 cm; stipules triangular, ca. 5 mm long, ca. 3 mm wide at base, stri-
ate, lately caducous, horizontal to reflexed; petioles striate, sparsely strigose,
up to 10 cm long, sometimes longer than leaflets, rachis up to 3.5 cm long;
stipels ovate to lanceolate, ca. 5 mm long; leaflets entire, terminal ones,
ovate to widely ovate, slightly oblique (lateral leaflets) at base, acute or acuminate
at apex, apiculate, 8-15 cm long, ca. 10 cm wide, the lateral leaflets some-
times with domatia within its basal veins, sparsely strigose on both sur-
face. Inflorescences in axillary pseudoracemes, glabrate or covered with uncinulated
or appressed, antrorse hairs, 10-50 cm long, with 10-40 flowering nodes;
buds 1—3 in each fascicle, 2 buds commonly flowering; primary bracts nar-

RAMIREZ AND DELGADO, A new Phaseolus species from Mexico 639

Fic. 1. Phaseolus albescens McVaugh ex R. Ramirez & A. Delgado, sp. nov. A. Portion of a
branch with leaves. B. I | from fresh material. C. Corolla (dissected), showing
a standard, two wings, and the keel with two tight coils at the apex. D. Androecium. E.
Gynoecium. F. Vexillary stamen. G. Flower bud. H, Calyx, dissected. I, Bracts and bracteoles.
J. View of the base (amplified) of the terminal leaflet lower surface. K. Pod. L. Seed. M.
Root. A—J, Machuca N. 4708; K—M, Ramtvez-Delegadillo 2537.

—

640 Sipa 184

rowly triangular to lanceolate, striate, persistent, 3—5 mm long, ca. | mm
wide at base; secondary bracts narrowly triangular, horizontal or reflexed,
ca. 2 mm long, | mm wide, subpersistent; pedicels thin, thicker at fruit-
ing, 1.5—2 cm long; bracteoles shorter than the calyx, lanceolate, never fal-
cate, slightly auricled at the base, 1—5-nerved, subpersistent, 3-4.5 mm
long, ca. | mm wide; flowers ca. 2 cm long, with ca/yx obliquely campanu-
late, sometimes with a bulged upper base, tube sparsely strigillose on the
outer and the inner surfaces, 5-6 mm long, upper lip emarginate, lower lip
with 3 subequal, triangular lobes, ca. | mm long; coro//a lilac fading to white,
and later ” ae standard oblong to orbicular, often little wider than
long, 1-1.4 cm long, 0.9-1.2 cm wide, distal portion of outer surface se-

‘cD.

—_

tose, ne with appressed, minute hairs, on the inner surface the lamina
shows a thickening at the point of reflexion, the surface between the bend-
ing point and the claw covered with micro-papillae, with two swollen ap-
pendages at each side of this basal portion, claw ca. 2 mm long; wings obo-

vate, ca. 2cm long, ca. | cm wide, constricted toward the base, upper basal
margin round-auriculed with a lamellate surface, claw of wing ca. 5 mm
long; keel ca. 1.5 cm long, claws 3 mm long, on distally 1.5 coiled, diam-
eter ca. 3 mm; staminal tube biauriculed towards the base, with sub-basifixed
anthers, vexillary or free stamen ca. 1.5 cm long, witha trapezoid-shaped
appendage toward the base, its edge undulated; pollen tricolporate, brevicolpate,
edge of the colpi reinforced by a margo, ectoaperture membrane granulated,
endoaperture (on non-acetolyzed pollen) covered with a smooth operculum;
exine with a distinctive sculpturing between the mesocolpium (foveolated-
—9); ovaries with 4—

pisilated) and the apocolpium (rugulated)(Figs. 4-5, 8
5 ovules, strigose, pollen brush short and laterally-placed on the last coil of
the style; st7gma apical, sometimes slightly introrse. Pods linear, slightly curved,

—9 cm long, ca. 1.2 cm wide, short tapered to tapered at base, and with a
ca. | cm straight beak at the apex; valves chartaceous with narrowly thick-
ened sutures, dehiscent, witha slightly glossy epicarp, ight brown-yellowish
coloured at maturity, sometimes purple-pigmented, striated, glabrate, (3)
4—S seeded. Seeds 1-1.3 cm long, 7-8 mm wide, ca. 3 mm thick (weight of

100 seed, ca. 20 ean reniform to sometimes quadrangular, compressed;
testa glossy and tan to d os brown mottled and streaked; hilum ovate with
a persistent ephilum, ca. 3 mm long, ca. 1.5 mm wide, rim aril prominent,
micropylar area lighter than testa color; lens prominent, slightly divided
in two. Seedling with epigeal germination; epicotyl pubescent; stipules bifid,
petiole with basal and apical pulvini (3-portioned); eophylls simple, ovate
with a cordate-truncate base, next leaves trifoliolate. Chromosome num-

aver: Ramirez-Delgadillo 2553 GBUG) (P.

ber, 2” = 22, in root-tip cells, vouc
Mercado-Ruaro, pers. comm.).

RAMIREZ AND DELGADO, A new Phaseolus species from Mexico 641

PARATYPES. MEXICO. Colima: Municipio de Colima. Rancho El Hee aos a 20
km 7 N de la Cd. de Colima, cerca de la Hacienda San Antonio, 19° 26' N; 103° 40! W,
1,306 7 Feb 1992, Rico 921 y Martinez (MEXU). Guerrero: Municipio « te i Union,
73 . . NE de Zilaacaneo carretera Zihuatanejo-Ciudad Alramirano, 1,77 2 Feb

1983, Tenorio et al. 3237 (MEXU). Jalisco: ee ih ane Sierra de Caer
(15-20 mi SE of Autlén) on the bajada south and west of the divide between Aserradero
San Miguel and Durazno, 2,000—2,350 m, 6 Nov | ion McVaugh 13943 (MEXU; MICH);
oe ~ Miguel, 1.5 km al E de su desembocadura, ca. 1 km al S de Rincén de Manantlan,

19° 3 ; 104° 12' 30 W, Sierra de Manatlan Ocidental, ae 800 m, 5 Jan 1985
Ee cz et al. 5113 (MEXU; WIS); Sierra de Manantlan, en el Rancho El Lamial, carretera
para el Rancho de woes 2,050 m, 10 Dic 1982, Calzada 9466 y ee (MEXU;XAL);
Municipio de Jocotepec, Barranca del Agua, al N de Zapotitdn, 18 Dic 1994, Machuca N.
4708 (IBUG),; aes Delgadilla 2553, 3600 ( (IBUG); Municipio de Tecalitlan, Sierra del
Halo, primitive road to San Isidro above the junction SSW of Tecalitlén, 2,000—2,200 m,
20 Nov 1959, McVaugh & aa 1240 Ie) 20 km SE of JCT with Route 110, on the
way to ae an via San Isidro, 2,130 m, 3 Dec 1995, Kajita et al. 95 120310 (MEXU).

n: Municipio de Coalcoman, W of Aguililla, 12 km SE of Aserradero Dos Aguas,
1,6( oe )O m, 27 Nov 1970, McVaugh 24754 oe MICH).

Distribution, habitat, and phenology —P haseolus albescens is restricted to western
Mexico, where the genus Phaseolus is well-represented (Del gado-Salinas 1985).
Sparse populations of P. a/bescens occur mainly in montane forests of the Sierra
Madre del Sur (1.e., Sierra de Manantlan and Sierra del Halo). Also on the
isolated mountain range known as the Sierra del Madrofio, located in the
center of the state of Jalisco, with geological links with the Trans-Mexican
Volcanic Belt. In Colima, Michoacan, and Guerrero it is distributed on mountains
of the Sierra Madre del Sur. The species grows in pine, pine-oak, and de-
ciduous forests, on brown and light-brown clay soils, always in humid and
protected environments at 1,300—2,100 m.

Phaseolus albescens its a late-fall and early winter bloomer (Oct. to Jan.),
and sets pods from November to March. In cultivation on the Universidad
de Guadalajara grounds, P. a/bescens grew to be a large and vigorous vine,
climbing up to ca. 10 meters high. It produced leaves every year and all
year around, flowering profusely from October to December, when X y/ocopa
bees were observed visiting its flowers. However, few fruits were set by this
plant each year. The plant lived for six years, until a severe drought killed it.

Common name and uses.—W hile people in Colima are aware of these plants,
which they called ‘frijolillo’, they do not eat them.

Wood anatomy of Phaseolus albescens (Figs. 2—3).—Wood is diffuse porous,
with shape of pores oval to angular, in distribution of radial multiples of 2—
10 vessels, aggregated in multiples chains or clusters; solitary vessels are of
uncommon occurrence. Numerous narrow vessels with 61 pores per mm?
mixed with the fewer wider vessels with 10 pores per mm?. Vessel elements
are short, with length that ranges from 174 to 192 pm. There is a consid-
erable range of variation, however, in vessel diameter, ranging from a mean

Sipa 18(3)

ae

ad ong
e
N
eS
4
-

4 6
lf

>
eos
jC ae |

a
~*,- pu

6 Sy
1D: a Sas
- a. ~~ =
=O@f. ==

A pene.
+

Fics. 2-3. Anatomical features of the stem of Phaseolus albescens. Fig. 2. Cross-section (x 2.5)
showing the secondary xylem with a diffuse arran :

gement of its vessels; also the presence of
a sclerenchymatous ring of fibers adjacent to the primary xylem. Fig. 3. Tangential section
(x 10) showing radial wood parenchyma composed of homocellular rays, with the presence
of nodules (typical of this species), indicated by an arrow. Fibers are located around vessels,
excluding the parenchyma cells, which contain abundant polyhedron crystals. Figs. 2—3.
Delgado et al. 1705.

RAMIREZ AND DELGADO, A new Phaseolus species from Mexico 643

——]

Fics. 4-9, Pollen grains hay eee polyanthus. Figs. 4-5. Light microscopic
ea of P. albescens: Fig. 4 ng rugulate ornamentation on the apocolpium. Fig.

5 showing the brevicolpus v ne a margo surrounding the furrow. Figs. 6-7. Light micro-
scopic photographs of P. polyanthus: Fig. 6 showing the finely-reticulated ornamentation
on the apocolpium. Fig. 7 showing the short colpus. Figs. 8-9. Scanning electron micro-
graphs of acetolyzed pollen grains of P. a/bescens. Fig. 8 showing at equatorial view one of
the three short colpus. Fi ig 2 Colas covered witha finely granulated membrane, and with
a slightly ag pore present due to acetolysis process. Scale bars =
LO pm. Figs. 4-5. Mochans N. 4708. Figs. a Debouck & Soto 1608. Figs. 8-9, Delgado et

al, 1705

644 Stipa 18(3)

tangential diameter of 42 jm in the narrow vessels to a tangential diameter
of 198 jm in the wider vessels. Perforation plates are all simple, the ones
from wider vessels elements are predominately transversal, while chose of
narrower vessels are oblique. Intervessel and vessel-ray pitting is alternate,
with minute pits from 5 to 8 pm.

Axial wood parenchyma is abundant and apotracheal, with 2 to 3 cells
per segment. Polyhedron crystals frequently forming long chains are abun-
dant in parenchyma cells that surround strands of fibers.

rs wood parenchyma are composed of scanty homocellular rays, with

3 to 4 rays per mm”. Ray cells are usually square to upright, where nodules
can be observed in tangential section.

The average length of the libriform fibers ranges from 741—1,667 pm
(with a mean of 1,310 pm), with a tangential diameter of 25 jam. Fibers are
located around vessels, their secondary walls reinforce the wall of the ves-
sels, but also excluding the parenchyma cells.

— description (Figs. a —Tricolporate, semitectate, spheroidal, P=
(40-)5 1.3(-55) pm length; E= (40—)55(-60) pm width. P:E=1.05. Polar
view subcircular, (47.2—)53. ae )) pm. Exine thickness of 2.4—3 pm. Colpi,
brevicolpate, covered witha finely granulated membrane, (19.2—)29.8(—30.4)
X (16.8-)21.7(—28.0) am. Pori slightly lalongate, 5 pm in diameter, the
pore is covered with a conspicuous operculum (non-acetolyzed pollen grains).

Observations on the floral biology of Phaseolus albescens. —Flowers of this spe-
cies were observed in Jalisco (N 19 36' 59"; W 103 30! 54.7"; 1,640 m) in
mid November where high visitation rates were registered, with numerous
incrafloral movements made by bees (Nylocopa guatemalensis, Bombus pullatus,
Bombus sp., and Apis mellifera). Observations occurred between 9:00 am and
noon. The color of flowers in this population was lilac fading to white and
later to yellow. White flowers were ignored and visits were confined to the
lilac-colored flowers, whose standard petal was normally raised and not al-
most fully reflexed as happens in the white-yellowish ones. The color change
(lilac to white) is accompanied by a change in the position of the standard,

which the bees probably perceived, so they are able to distinguish between
the young and older flowers, apparently there is both an optical and posi-
tion change to aid the insect in its visit.

Although flowers of a wild plant of Phaseolus vulgaris were close-by, no
visitation of bees between both species were detected. Also, plants of wild
P. coccinens were seen growing within two hundred meters, no hybrids be-
tween them or with P. vv/garis were found.

Relationships with the wild P. polyanthus.—As noted in the introduction,
McVaugh (1987) recognized differences between plants corresponding to
Phaseolus albescens and plants of wild P. coccineas and cultivated plants of P.

RAMIREZ AND DELGADO, A new Phaseolus species from Mexico 645
polyanthus. Recently, the wild progenitor of the cultivated P. polyanthus has
been identified in Gutemala by Schmidt and Debouck (1991), alchough
suill not completely described, observations made on its general morphol-
ogy (Debouck & Soto 1608; WIS) and on its seeds (CIAT # 35877) show con-
sistent differences between the two.

Indeed, P. a/bescens is superficially very similar to the wild P. polyanthus in
fruit, but the number of seeds in the fruit of each taxon differ, mainly hav-
ing 4—5 seeds in P. albescens, while pods with 3—4 seeds are present in the
Guatemalan plants. Also the flowers of the two species differ in a number
of ways, most noticeably in the form and size of the bracteoles, which in P.
bolyanthus are longer and broadly lanceolate, in addition to the contrasting
floral coloration, and change in position of the P. a/bescens standard petal.
The pollen of P. a/bescens can be distinguished from P. polyanthus by larger
dimensions (P:E ratio > 0.13) and foveolate-rugulate exine ornamentation
instead of a finely-reticulated one. Furthermore, their actual disjunct dis-
tribution (P. albescens restricted to west-central Mexico and wild P. polyan-
thus mainly to Guatemala), apparently endures from a fairly old separation
from each other in the past. This can be assumed comparatively by the number

of molecular changes presented by P. alhescens different from the cultivated

and wild P. polyanthus.
The specific epithet ‘a/bescens’ refers to the flower color of this species being
lilac at anthesis, then turn in, white and later yellow when older.
KEY TO PHASEOLUS ALBESCENS AND WILD RELATIVES

|. Inflorescences few-flowered, commonly with 8 or less flowering nodes; flowers
small to medium size, standard-petal 1 cm or less long; pods narrow, ca. 1
cm wide; seedlings with epigeal germination.

2. Pedicels in flower 3-7 mm long; bracteoles linear-ovate to lanceolate, shorter

than calyx, less than | mm wide, 1—3 veined; petioles of first seedling leaves

(eophylls) subsessile P. acutifolius A. Gray

Pedicels in flower 0.5-1.2 cm long; bracteoles broadly ovate, prominent,

equaling the calyx, 2.5-4 mm wide, 5—10 veined; petioles of first seed-
ng leaves (eophylis) elongated, not sessile

L. Inflorescences many-flowerec

2)

P. vulgaris L.
,commonly with 10-30 flowering nodes; flowers
medium to large size, standard-petal more than | cm long; pods broader,

more than 1 cm wide; seedlings with epigeal or hypogeal germination,

3. Plants developing of a thick, often branched, tuberous root: corolla com-
monly red, rarely purple or white; stigma apical-extrorse; seedlings with
hypogeal germination

P. coccineus L.
3. Plants not developing of a tuberous root, main root lignified or fibrous;
corolla dark pink, purplish, lilac or white; stigma terminal or introrse;
seedlings with epigeal germination.
4. Corollas dark pink or lilac to purple, wing-blades at anthesis not fully
expanded, clasping; bracteoles broad, 2.5—-8 mm wide; plants from Costa
Rica and Panama

P costaricensis Freytag & Debouck

646 Stpa 18(3)

4. Corollas lilac-white or white, wing-blades at anthesis fully expanded;
bracteoles narrower, 1—1.5 mm wide; plants from western México and
Guatemala.

5. Bracteoles broadly lanceolate, sometimes falcate, commonly exceeding

the calyx, 6-8 mm long; pods with 3-4 seeds .......... P. polyanthus Greenman
5. Bracteoles lanceolate, never falcate, shorter than calyx, 3-4.5 mm
long; pods with 4—5 seeds. Phaseolus albescens

McVaugh ex R. Ramirez & A. Delgado
ACKNOWLEDGMENTS

We thank Calixto Le6n-Gomez for his invaluable help in the anatomical
work and P. Mercado for providing us with the chromosome count. We also
thank Ricardo Ayala for the identifications of the bees, Noemi Jiménez Reyes
for the pollen photographs under light microscopy, and Sara Fuentes for
providing the SEM photos of the pollen grains. We extend our gratitude to
A. Wong, F. Basurto, and Gabriel Flores for their help in the field, and to
M. R. Garcia Petia (MEXU) for kindly requesting loans for this study, and
to the Curators of the following herbaria: IBUG, MEXU, MICH, US, WIS,
and XAL. The seeds used for comparison with wild Phaseolus polyanthus were
provided by CIAT. Thanks are also due to Dr. Fernando Chiang for com-
posing the Latin diagnosis and to an anonymous reviewer for useful criti-
cism. Profesora Ma. del Refugio Vazquez Velasco prepared the illustration

of P. albescens.

on)

REFERENCES

DELGADO-SALINAS, A. 1985. Systematics of the genus oe (Le aera in North and
Central America. Ph.D. dissertation. University of Texas-Aust 545.
DELGADO-SALINAS, A. oe Variation, taxonomy, c oe. ie germplasm poten-
tialities of Phaseolus coccinens L. In: P. Gepts, ed. Genetic resources, domestication and evo-
lution of Phaseolus xo Kluwer Academic Publishers, Dordrecht, Holland. Pp. 441—466.
DELGADO-SALINAS, A., oe RLEY, A. RicuiMANn, and M. Lavin. 1999. Phyl a analysis
f the calewated ae Gia species of Phatcoles ies: Syst. Bor. 24(3). I
McVa iGH, R. 1987. Phaseolus. In: W.R. Anderson, ed. 0 eee Galiciana. Vol. ise ese
The University of Michigan Press, Ann Arbor, U.S.A. Pp. 654-655.
RAMIREZ- DE LGADILLO, R. me ‘Taxonomia, at gta y erat del género Phaseolus
e Tlajomulco de a ¥ Jocotepec, Jalisco. Tesis de

1.

los muni

ence Agrénomo, Unive od de Guadalajara, Méxi
Scumit, V. and D.G. Desouck. 1991. Observations on ay origin of Phaseolus polyanthus
Greenman. Econ. Bot. 45:345—364.

THE RELATIONSHIPS OF THE AMERICAN
BLACK-FRUITED HAWTHORNS
CRATAEGUS ERYTHROPODA, C. RIVULARIS
C. SALIGNA AND C. BRACHYACANTHA
TO C. SER. DOUGLASIANAE (ROSACEAE)

J.B. PHIPPS
Department of Plant Sciences
The University of Western Ontairo
London, Ontario, NOA 5B7, CANADA

>

ABSTRACT

A group of black-fruited Crataegus (Rosaceae) from the western United States is analysed.
Crataegus erythopoda, a species with previously poorly unde erstood affinities, is shown to ¥
closely related to C. rivularis. The affinity of C. rivalaris and C. saligna is reconfirmed. T
three species form a clade subtended by C. brachyacantha and different from the clade con-
taining C. douglas and C. okennonii when analyzed by PAUP using 38 morphological char-
acteristics. Crataegus rivularts and C. saligna are lec one

RESUMEN

Se estudia un grupo de Crataegus (Rosaceae) con frutos oe del oeste de los Estados
Unidos. mead erythopoda, una oe con afinidades pobremente conocidas

, se muestra
que esta y relacionado con C. rivilaris. Se reconfirma que C. rivwlaris es también muy
afinaC. ees Las tres ae en un clado en cuya base se encuentra C. brachyacantha
y que es diferente del clado que contiene a C. douglasii y G kemanit a se analiza
mediante el PAUP usando 38 caracteristicas morfoldgicas. Se tipifican Crataceus rivularis
y C. saligna.

Black-fruited Crataegus are in a substantial minority worldwide, as also
in North America where perhaps seven species are black-fruited. Numer-
ous authors have pointed out the significance of fruit color in relation to
frugivor dispersal and this topic has even received a little attention for North
American hawthorns (e.g. Sallabanks 1993). It is not known whether North
American black-fruited hawthorns form a natural clade and without un-
dertaking an exhaustive examination that potentially includes many red-
fruited species, this fact cannot be determined. However, it had appeared
to me that they fell into at least two groups, the first being all members of
series Douglasianae (Rehd. ex Sarg.) Rehd. (this includes C. douglasii Lind|.,

C. suksdorfir Sarg.) Kruschke and C. okennonii J.B. Phipps) together with a
second group (C. brachyacantha Sarg. & Engelm., C. erythropoda Greene, C.
rivularis Nutt. ex Torr. & A. Gray and C. saligna Greene) which might have

Sipa 18(3): 647-660. 1999

648 Sipa 18(3)

some loose interrelationship. I test chis hypothesis of two clades with a cla-
distic analysis using 38 morphological characteristics.

Series Douglasianae has been the subject of a substantial amount of re-
cent research and is more northerly than the group of species on which |
focus in this paper, with a southern limit of about 43° N except in Califor-
nia where it is farther south. Love and Feigen (1978) showed that C. douglasi
could hybridize with the distantly related and introduced C. monogyna Jacq.,
Brunsfeld and Johnston (1990) provided solid grounds for raising C. swksdorfir
(Sarg.) Kruschke from a variety of C. douglasii to specific rank while Dickinson
et al. (1996) contrasted the breeding systems in the 20-stamen C, swksdorfiz
and the 10-stamen C. douglasii to good effect. Dickinson and students are
continuing their biosystematic and morphometric analyses of ser. Douglasianae.
Recently, I described a new species, C. okennonii, from this group (Phipps &
O’Kennon 1998). Due to this recent activity and the continuing researches
of the Dickinson group on ser. Douglasianae, | restrict coverage of the series
in this paper to the cladistic analysis merely to discover if ser. Dowglasianae
is a separate clade from the other species considered.

Here, therefore, I direct attention to the more southerly, and allopatric,
— of the ‘black-fruited’ Crataeg?, the group of C. rivilaris, C. erythropoda,

C. saligna and C. brachyacantha. The relationships of these species has al-
ways been much more controversial. Crataegus saligna was placed with C.
brachyacantha by Palmer (1925), a supposition that I followed without study
in the Maloid checklist (Phipps, et al. 1990). Then, particularly influenced
by field observations, I recognized its probable affinity to C. rzvilaris, a re-
lationship in fact explicitly noted by Greene (1896) when he described C.
saligna. With regard to C. erythropoda, as recently as this year (Phipps 1999),
I included this species among the western red-fruited species. Neverthe-
less, the existence of a few intermediate specimens noted since that paper was
prepared, together with the detailed examination for this paper, and prepa-
ration of the draft description for the Flora of North America, showed con-
clusively that it was actually close to C. rivularis. Also recently, Welsh (1982)
placed both C. rivularis and C. saligna under C. douglasi1, a solution that I do
not believe that any current student of Crataegus would consider tenable

I therefore resolved to investigate the cladistic interrelationships of the
North American black-fruited species and, in the cases of C. saligna, C. rivularts
and C. erythropoda, which have never received any independent taxonomic
study since their original descriptions, in contrast to ser. Dowglasianae, to
carefully characterize and typify them. In addition, C. rivalaris and C. saligna
are provided with detailed distribution maps for the first cime. Finally, I
need to comment on my cautious ‘black fruited’. Species in this group either
have black, purple, or burgundy fruit when fully ripe, the particular color,

Puipps, Black-fruited Crataegus from the western United States 649

and its changes during ripening, having taxonomic significance. The term
‘black-fruited’ in the title therefore refers to a group of related hawthorns
that are predominantly, but not entirely, black-fruited at full maturity.

METHODS

An extensive sample of herbarium specimens of the taxa to be studied
has been assembled by my own fieldwork in recent years, assisted by R.J.
O’Kennon, together with many examples of loan specimens. As this paper
is not aimed at assessing specific limits, no morphometric phenetic analy-
sis of large samples was conducted. However, the samples were used to de-
termine the characteristics of synthetic OTU’s, one for each species, that
would be analyzed by PAUP 2.4.1. The 38 characteristics used for this purpose
are listed in Table 1. Distribution maps for previously unmapped or inad-
equately mapped species were created by Range-Mapper, a program cre-
ated by the firm Tundra Vole of Fairbanks, Alaska. For this purpose, where
sufficiently precise location data existed on the herbarium label this was
converted into latitude/longitude coordinates accurate to the nearest minute.
Files for each species were then mapped. Typification followed standard
procedures. Syntypes or potential lectotypes were located, occasionally with
great difficulty, and hololectotypes or neotypes selected on the basis of goodness
of fit to the protologue, citation in the protologue (if pertinent), and speci-
men quality, when a choice was available. No specimens are illustrated due
to the existence of excellent illustrations elsewhere. The characterizations
presented here of the species outside ser. Dowglasianae are in fact somewhat
detailed diagnoses because this work does not purport to be a taxonomic
revision, because excellent descriptions exist elsewhere and because, in my
view, they represent four well-marked species even by conservative criteria.

—_—

INTERRELATIONSHIPS

The possible and intriguing relationship between C. saligna and C. rivularis
has already been suggested. If these two proved to be sister species then a
new series ‘Rzvulares’ could be created to accomodate them. However, on
reflection, it seemed that C. rivlaris might be even closer to C. erythropoda
on the basis of identical floral and inflorescence characteristics and fruit differing
only in fully ripe color. A series that included both C. saligna and C. erythropoda
is perhaps a little broader than most series in North American Crataegus.
We need also to pay attention to Palmer’s (1925) view that C. saligna was
closely related to C. brachyacantha and therefore should be placed in ser. Brevispinae
(Palmer used the nomen nudum Brachyacanthae), where I had provisionally
placed it (Phipps et al. 1990) following convention and without study. Crataegus
brachyacantha is very similar to C. saligna in flower and leaf characteristics

650 Sipa 18(3)

Taste |. Thirty-eight morphological characteristics scored for cladistic analysis.
Plant, general 21. Leaf teeth: glands

Inflorescence
22. Flower: number
23. Pedicel: cad
Thorns and Twie 24. Anchesis cit
1. Thorns: Peace present? eee
a OWE)
gh 25. Diamete

26. Calyx ee margins
)

ak a no.
Ean ie 28. Anther: color
9. Twigs, | yr old, sie browns 3
LO. Twigs, 1 yr old, color; gray

1 Bruit
ad 7 :
af 30. Fruit: color, month before full ripeness
I 1. Lamina: length : ins
31. Fruit: color at full ripeness

12. Lamina: length/breadth Tae
, ; 32. Fruit: shape, |
13. Lamina: position of widest part a Teer :
| 33. Fruit: shape, 2
14. Lamina: venation, number be ee
j {. Fruit: pubescence
15. Lamina: veins to sinuses - ae ; .
. 35. Fruit: calyx orientation
16. Lamina: lobe number , eae :
36. Pruic: length (height)
de shape eae :

>

17. Lamina: lob

~)

: Pyrenes: lateral faces
18. Lamina: max. sinus depth (LID
19. Lamina: abaxial pubescence, young ap ae foliage
20. Petiole: glands olor
§

but differs greatly in the thorns (curved, < 1.5 cm long), color of overmature

y

inflorescence (yellowish orange), fruit size (generally larger), smooth lat-
eral faces of the pyrenes, and lack of copper-colored bark. Also Welsh had
made both C. sa/igna and C, rivilaris varieties of C. douglasi1, in what seemed
to me to be an untenable association. It therefore seemed pointful to con-
duct a numerical taxonomic study to throw light on these contrasting pos-
sibilities and to establish whether any of these taxa were especially close to
members of ser. Dowglasianae.

For this, | scored 38 characters (Table 1) for 9 synthetic OTUs, one for
each species, which included all the black-fruited Crataegis species that are
the particular subjects of this paper (C. sa/igna, C. rivularis, C. erythropoda
and C. brachyacantha) plus C. douglasii 5.8. , C. suksdorfii and C. okennonii of
ser. Douglasianae as well as the red-fruited outgroups C. mo//is (Torr. & A.Gr.)
Scheele and C. monogyna Jacq.

Using PAUP 2.4.1, C. erythropoda and C. rivilaris are shown to be sister
species in the three shortest trees (Fig. 1). The dovglasi7 group, consisting
oe douglasii and okennoni1, always formed one sub-clade while C. brachyacantha,

O. saligna, C. rivilaris and C. erythropoda always formed another in all shortest
oe However, C. saligna and C. brachyacantha were by no means
very close in these cladograms although C. rivularis and C. erythropoda were

Puipps, Black-fruited Crataegus from the western United States 651

monogyna PAUP 2.4.1
douglasii 9 OTUs

38 characters

okennonii

brachyacantha

erythropoda

rivularis

saligna

mollis

|. Cladogram of North-American black-fruited Crataegi created by PAUP using 38
ao characters.

shown in all analyses to be sister species. The two unpublished trees only
differ in the location of C. saksdorfii which is either basal to C. douglasii /
okennonii (published tree), basal to (B(Sa(R,E))), using their epithet abbre-
Viations, in one unpublished tree or basal to both (D,O) and (B(Sa(R,E)) in
the other.

The union of all seven black-fruited species inside the red-fruited out-
groups C. mollis and C. monogyna does not, however, indicate that the black-
fruited group is monophyletic. An analysis using all the red-fruited species
would be required to generate such a finding and that is not the function of
this exercise. Rather, it is to locate the position of C. erythropoda, C. rivularis,
C. saligna and C. brachyacantha relative to the immediate C. douglasii group.
The cladogram endorses the view that neither C. sa/igna nor C. rivularis are
part of the species C. douglasii. The following key summarizes the differ-
ences among the four species in rivu/aris clade.

KEY TO RIVULARIS-CLADE OF BLACK AND BURGUNDY-FRUITED CRATAEGUS

l. oe 20; anthers cream; flowers 10-12 mm diam.

2. Thorns |—1.5 cm long, decurved, grayish; twigs grayish; c flowers
orange-yellow; nutlets with lateral faces plane C. brachyacantha
2. Thorns 2—4 cm long, straight, black; twigs copper-colored; overmature
flowers white to dirty white; nuclets with lateral faces pitte C. saligna
1. Stamens 10; anthers pink; fowers 15—17 mm diam
3. Leaves essentially aes on more than 2 times as long as broad,

widest near the middle y ripe fruit blac C. rivularis

3. Leaves evidently ate 3- me d per side; usually ca. 1.5 times as long as
broad, usually widest in the basal third; fully ripe fruit usually burgundy
. erythropoda

652 Sipa 18(3)
TAXONOMIC AND NOMENCLATURAL NOTES ON INDIVIDUAL SPECIES

CRATAEGUS ERY THROPODA—Characterization
Crataegus erythropoda Ashe, North Carolina Agric. Exp. Sta. Bull. 175:113. 1900.

Crataegus bakeri Greene, nomen nudum

This species has been described, illustrated and mapped in my recent paper
(Phipps 1999) where it was entered into a new monotypic series, Cerrones
J.B. Phipps, on account of its distinctness from other red-fruited species.
Crataegus erythropoda may be briefly characterized by its ovate-rhombic, +
glabrous, shallowly-lobed, smallish leaves; glabrous inflorescences with LO-
stamen flowers having pink-purple anthers; suborbicular fruit which is burgundy
at maturity (I have never once seen ‘orange’ as described in the protologue
by Ashe (1900) and Holmgren (1997)); shiny coppery bark on 2—5 cm thick
stems and thorns mainly 2—4 cm long. Crataegus erythropoda occurs in sites
with groundwater available, or otherwise mesic, mainly in intermountain
Colorado and adjacent New Mexico, where it is quite common. Holmgren
(1997) also records it from eastern Utah and northeastern Arizona. The ‘or-
ange’ fruit color cited may refer to immature fruit since the type was col-
lected in August, but I have not myself seen this color even in mid-Au-
gust. Crataegus erythropoda has presented no serious problems in delimitation
or recognition though its relationships have been hitherto obscure. See the
procologue (Ashe 1900) for another full description. Interestingly, Palmer
(1925) assigned C. erythopoda to ser. Douglasianae, an idea taken up by no-
one else, but nevertheless the closest approximation to date. Observations
made for this paper indicate that at anthesis C. erythropoda is almost indis-
tinguishable from C. rivu/aris in floral/inflorescence characteristics. Significant
differences in leaf shape and ripe fruit color exist, however, as indicated in

the key.

A specimen of Greene's from the Lower Cimarron River, Colorado (NDG),
collected in 1896 and labelled ‘C. baker7’, a name never published, is actu-
ally a perfectly adequate specimen of C. erythropoda Ashe.

CRATAEGUS RIVULARIS—Synonymy, Characterization and Typification
Crataegus rivularis Nutt. ap. Torr. & A. Gray, Fl. N. Amer. 1:464. 1840.
Crataegus douglasii Lindl. var. rivularis (Nutt. ap. Torr. & A. Gray) Sarg., Gard. &
Forest 4:81. 1902
As with C. erythropoda, there have not been serious problems in taxon
recognition or delimitation although some floristic authors have followed
Sargent (1889) in relegating C. rivwfaris co varietal rank under C. douglass.
Crataegus rivularis is a locally common, even locally abundant, species of
intermountain USA, found in many locations in this generally dry area where
there is a high water-table. I map this widespread species (Fig. 2) which

Puipps, Black-fruited Crataegus from the western United States 653

occurs from southern Idaho to northern New Mexico and from south-cen-
tral Utah east to the Rocky Mountains. It reaches 6—7 m in favorable sites
where it may form extensive thickets. The bark of 2 cm diameter branches
is copper-colored like C. erythropoda and the generally fine thorns are mainly
2.5-4cm long. The normally elliptic leaves are tapered at each end, subglabrous,
4—6-veined and quite unlobed on short-shoots, being beset with many sharp,
forward-pointing fine marginal teeth. The glabrous inflorescence bears large
10-stamened flowers with pink anthers and can hardly be differentiated from
that of C. erythropoda. The numerous, often large, suborbicular fruit pass
through a striking deep reddish-burgundy color to deep purple and finally
black. See also Sargent (1890) for a detailed characterization and excellent
plate by Faxon. Intermediates with C. douglasii are not known and C. rivularis
is essentially allopatric with that more northern species which facts rein-
force the findings of the cladistic analysis. However, because Welsh (1982),
Holmgren (1997), and Sargent (1889, 1890) have included C. rivularis in
C. douglasii, | would like to emphasize their distinction with the following
key couplet.

1. Thorns fine, little recurved, usually 2.5—5 cm long; bark of 1-2 cm diam.
branches shiny coppery brown; flowers 14-18 mm diameter, calyx lobes 5—
8 mm long, long-attenuate from a broad base; leaves narrow elliptic, unlobed
or rarely with |—2 very small lobes per side, tapered at both ends, acute to
acuminate at the apex, the venation semi-camptodromous; fruit subglobose,
11-14 mm diam., crimson-lake ripening to shiny black >. rivularis

. Thorns stout, often recurved, usually 1.5—3 cm long; bark of 1-2 cm diam.
branches tan to gray-brown; flowers 12-15 mm diam., calyx lobes 3-4 mm
long, narrow-triangular from a piped base; leaves much broader, most com-

monly broad elliptic to narrow-obovate in general pode usually | ee alepes
7

nar n

in rare narrow-leaved specimens), the
in rare narrow- eee specimens ane obtuse to Spar ate at the apex; fete

generally ellipsoid, usually S 9 mm diameter, ripening dull purple to oe
black or black, generally with rather strong bloom C. douglasii, s.s.

The suite of differences is so large, together with the lack of intermediates
and different distribution, that one cannot reasonably place C. rivularis in
the same species as C. dowglasi?.

In making C. rivularis a variety of C. douglasii Sargent’s overall under-
standing of this taxon was poor, illustrated by his providing (1890) an in-
accurate range extension from northwestern California to Puget Sound and
his statement, “usually a low intricately branched.....shrub.” This perhaps
helps to explain the cautious rank chosen by Sargent, a notorious splitter
by modern criteria.

Crataegus rivularis has not been typified, therefore, I lectotypify it here.
The type description for C. rivularis (Torrey & Gray 1843) is of characteris-
tic brevity and imprecision for the period and might at first sight refer to

654 Sipa 18()

eis Se 5: an -103°

| = :
I —T
I ‘
, . _ NB
| a \ -
l oe
| | a =
|
|
; | a
| =. CO
I a
I =
I
37° I'UT I
he
\
ae I
\ !
|
NM !
a |
|
I

Fic. 2. Distribution of C. rvvwlaris Nutt. ap. Torrey and A. Gray from collated herbarium
records.

C. douglasii or even C. saligna, as well as to the taxon normally associated
with the name.

The protologue for C. rivilaris cites “Oregon, rivulets in the Rocky Mountains”
as the type area with Nuttall as collector. It is hard to know exactly where
this is, because of the extensive nature of che Oregon Territory in 1843.
What we call C. rivalaris today is a distinct, well-collected taxon with a
well-established range (Fig. 2) which just reaches southern Idaho, part of
which is an acceptable area for “Rocky Mountains.” Putative syntype ma-

Puipps, Black-fruited Crataegus from the western United States 655

terials of C. rivalaris from BM, GH and PH were borrowed in order to lectotypify
this species.

Of the two PH specimens sent (both on the same sheet) one (on the right-
hand side) can be rejected directly as it is a Canby-collected C. douglasii
from 1873, while the left-hand specimen, labelled ‘Rocky Mountains’, is a
leafy twig without reproductive material that requires comparison with the
protologue. Both the BM and GH sheets have specimens collected by Nuttall
that are a reasonable match for the somewhat poor description in the protologue
and are clearly the same species as each other. In view of the indifferent quality
of the protologue and the consequence that a species other than what is con-
ventionally called C. rivu/aris might have been described, I provide in Table
2 critical comparisons between the protologue and putative types (cols. 1,
2), between the protologue and C. douglasii (col. 3) and between the protologue
and standard interpretation of C. rivilaris (col. 4).

Several points in the protologue cannot be assessed from the putative syntypes
from BM and GH. These are the arborescent nature of the plant (not re-
corded on the collection label) and flowering characteristics (the putative
syntypes are post-flowering). Also, Nuttall in describing the leaves as “ovate,
obovate” poses something of a problem, as these are not terms that we would
apply today to the putative syntype leaves, which are clearly nearer to nar-

rowly elliptic. In fact, in examining many protologues for C. series Coccineae
drawn up by C.S. Sargent in the first and second decades of this century, |
have observed that “ovate” is almost a generic leaf-shape. In my opinion,
having also examined many other Crataegus type descriptions from the nineteenth
century and earlier, the term “ovate” was not used with its present preci-
sion and therefore, with regard to leaf-shape, the protologue may be said to
describe the material of the putative syntypes in question with adequate
accuracy for the period. The other characters match the protologue. As to
the PH putative syntype (col. 2), the matches between it and the protologue
are so few as to be almost meaningless. Moreover, the specimen possibly
belongs to C. erus-galli, as previously stated by Eggleston in an annotation
on the sheet, but this is a species never found in the Rocky Mountain area.
However, the specimen also resembles C. sa/igna, but because it is sterile,
conspecificity is uncertain. The final two columns in the table compare the
protologue with typical C. dowglasii and what is normally called C. rivalaris
(alternatively C. douglasii var. rivilaris). Because C. douglasii has lobed leaf-
blades and “short” thorns, whereas C. rivi/aris has longer thorns and unlobed
leaves, the latter constitutes the better match for the protologue. Thus, the
choice of lectotype lies between inadequate vegetative material of what is
possibly C. saligna at PH and adequate specimens clearly representing what
is normally called C. rivwlaris at BM and GH. Therefore, I lectotypify

—_—

656 Sipa 18)

3LE 2. C ompar ison of protolog ue characteristics with putative syntypes of C. vivilaris (see text) and
with i. of C. ae and C. rivularis.

match for match for
putative putative match match
syntypes syntype for C. for C.
protologue characteristics at BM,GH at PH douglasii rivularis
Arborescent ? ? + :
Nearly glabrous + ' + '
Not glandular + i + 4
Leaves - ovate or obovate 2? ?? +,4 2?
- obtuse or sometimes acute . + t
- simply or somewhat incisely + ‘ * ,
serrate
- attenuate into a short petiole : t +
Thorns - “spines” long + ? ° A
Corymb - many flowered ? ? + '
- glabrous ? ? us ;
- flowers small ? + ;
- segments of calyx obtuse and + ? 4 !
very short
Fruit - black ? ? + ;

- cannot determine a match

+ = perfect match

+ = reasonable match
us = usually a match

° - not a match

C. rivilaris with the Nuttall specimen “Rocky Mountains” (holotype BM;
isotype GH) and retain the usual interpretation of this taxon.

CRATAEGUS SALIGNA—Synonymy, Characterization and Typification
Crataegus saligna Greene, Pittonia 3:99. 1896.

. die wheeleri A. Nelson, Bot. Gaz. 34:369. 1902.
louglasii Lindl. var. duchesnensis S.L. Welsh, Great Basin Naturalist 42:9. 1982.

ee saligna isa medium-sized to large thicket-forming bush grow-
ing along streams and other locally moist areas in incermountain Colorado
and northeastern Utah (Fig. 3). Its range is the most restricted of the four
species dealt with in this paper. In many respects it is like C. r7vularis but
it has smaller leaves, thorns, flowers and fruit and 20 cream instead of 10
pink anthers. The fruit is more fully black (darker than C. rivalaris when
sub-ripe in lace August) and the leaf-teeth are quite different from C. rivularis,
being small, numerous and obtuse, rather than longer and sharp. Moreover

Puipps, Black-fruited Crataegus from the western United States 657

|

Fic. 3. Distribution of C. sa/igna Greene from collated herbarium records.

there are usually 6—9 lateral veins on one side of a leaf, compared to 4—5 in
the much larger leaf of C. rivu/aris. Indeed, the leaves are remarkably simi-
lar to those of C. brachyacantha. Crataegus ie has sometimes been con-
fused with C. douglasi from which it has even mor nees than C. rivilaris
and it was erroneously called C. dowglasii var. Dicbennate o Welsh (1982).
It is similar to C. rivularis in its coppery bark on 2—5 cm diameter branches,
and slender, + straight chorns usually 2—4 cm long [except recurved and
0.75-1.5 cm long in WW. Robbins 6972 from Newcastle, Colo. (COLO)},

narrow leaves, intermountain distribution and similar habitat.

Crataegus saligna was described by Greene (1896) from the Lower Cimarron
River, Colorado, without citation of type material, a situation that has not
been remedied to date. My search for potential lectotype material involved
the main Greene herbarium at Notre Dame University, Indiana (NDG),
WIS and NA where remnants of the remainder of Greene’s herbarium had
been dispersed from LCU, and CAS, COLO, NY, UC, and US. Only one
herbarium generated a putative lectotype, the specimen being E.L. Greene
s.m., Colo., Cimarron, 31 Aug 1896 (NY). I therefore lectotypify C. saligna
by this specimen. Fortunately, the type description of C. sa/igna is entirely
adequate for species recognition, such that this taxon has become a well-
accepted element in the Colorado flora (Harrington 1964; Weber & Wittmann

658 Sipa 18(3)

1992). Sargent (1902) provided a more extensive description of C. saligna,
together with the citation of specimens dating back to 1845 and a fine il-
lustration by Faxon.

Crandall’s 1896 collection of C. sa/igna (at RM) from the Lower Cimarron
River also became a potential lectotype for this species because the Lower
Cimarron River, Greene's type location for C. sa//gna, enters the Black Canyon
of the Gunnison. However, there is no indication that Crandall’s specimen
was ever studied by Cea and therefore it is rejected for this purpose.

Wheeler also collected several hawthorn specimens from the same area,
but in 1898, too late to be lectotype material of C. sa/igna but which it is
convenient to comment on here. From among these, Aven Nelson (1902)
described C. wheeler? which I lectotypify by H.N. Wheeler 523, Black Can-
yon of the Gunnison (RM). As this ts identical to C. sa/igna, C. wheeler’ becomes
a synonym of that species.

Complicating the taxonomic picture was $.L. Welsh’s description (Welsh
1982) of a new variety of C. donglasii, var. duchesnensis 8.L. Welsh, from northeastern
Utah. From the type description, though very brief, chis new taxon appeared
to me to resemble C. sa/igna rather then C. douglasi7. Loan of relevant ma-
terial from BRY proved this suspicion to be correct, consequently C. dowglasii
Lind]. var. duchesnensis $.L. Welsh is here synonymized under C. saligna. Also,
some of Welsh’s specimens annotated as his new ae of C. douglasii proved
to be C. rivularis and I therefore supply hereunder a list of Utah specimens
cited by Welsh (1982) as C. douglasit var. duchesnensis that I identify as C.

saligna:

UTAH: a Co.: 5 mi N of F ruitland, « = fe on J Brotherson 5O8;, 5 mi NW
of Hannah, 17 Aug 1965, K.S. Erdman 2522; 'T15 ROW Sec. 36, 9 J 9706, ae :
Hansen s.n., 24 km NW ¢ of Duchesne, along Utah 35, i Sep 1970, o L. Welsh, N
inte G. Moore 10928 (HOLOTYPE). Uintah Co.: Merkley Park, 18 May 1963, o
W ‘elsh GG. Moore 1948.

These records, and others of my own generated by this discovery, repre-
sent quite a significant range extension for C. sa/igna, a species hitherto known
only from intermountain Colorado. | therefore offer what I believe to be
the first published range map (Fig. 3) of C. saligna.

CRATAEGUS BRACHYACANTHA Sarg. & Engelm.—Characterization
Crataegus brachyacantha Sarg. & Engelm., Bot. Gaz. 7:1 1882.

Crataegus brachyacantha \ies among the select group of hawthorns with-
out synonymy, this fact alone attesting to its distinctiveness. It is one of the
largest North American hawthorns, occasionally pacing a 14 m tall, though
it is more usually a bush or small tree at maturity 4-8 m tall. It occurs
naturally throughout Louisiana and the adjacent parts of all surrounding
states reaching its best growth on moist bottomlands. The twigs are beset

Puipps, Black-fruited Crataegus from the western United States 659

with few to numerous very short (ca. 1 cm long) recurved thorns, unique in
Crataegus to my knowledge. The short-shoot leaves are elliptic, unlobed,
very glossy, glabrous with numerous secondary veins and crenate margins.
They resemble the leaves of C. sa/igna and color brilliantly in autumn. The
multi-flowered inflorescence is glabrous and turns orange-yellow when over-
mature, also apparently unique in Crataegus. The small flowers have 20 sta-
mens. The fruit, dead-black at full maturity, is covered by a dense bloom
before the bloom is abraded, and together with a more purple-black sub-
mature skin, the fruit may look ‘blue’, hence the vernacular name ‘blue-
berry haw’. The fruit is somewhat bitter and has five plane-sided nutlets.
Full descriptions and illustrations may be found in Phipps (1998) and Sargent
(1902). The species is also mapped in detail in Phipps (1998). A white-
fruited variant, unique to Crataegus in this respect, was seen in 1922

There is no problem in delimiting this species and it has never been thought
to be especially closely related to, or a part of, C. douglasii.

ACKNOWLEDGMENTS
I thank the curators of BM, BRY, CAS, COLO, GH, NDG, NY, PH,
RM, UC, US, and WIS for loans of specimens and/or cooperation in the
search for lectotype material. I also thank the National Sciences and Engi-
neering Research Council of Canada for funding that supported this work
and an anonymous reviewer for the suggestion that NY might possess a
potential lectotype of C. sa/7gna, a circumstance that prevented neotypification.

REFERENCES
AsHE, WW. 1900. Crataegus le North Carolina Agric. Exp. Sta. Bull. 175:113.
ee, S.J. and F.D. JOHNSON. 1990. Cytological, morphological and p oo support
for specific status of Crataegus ane (Rosaceae). Madrofio 37:274-28
DICKINSON, T.A., S. BELAOussorF, R.M. Love, and M. Muntyamma. 1996. North American
black-fruited hawthorns (Crataegus sect . Douglasii Loud.): 1. Variation in floral construction,

breeding system correlates anne their possible evolutionary significance. Folia Geobot.
Phytotax. 31: 355-371.
GREENE, E.L. 1896. Crataegus saligna n. sp. Pittonia 3:99.
HarrincTron, H.D. 1964. Manual of the plants of Colorado - a biodiversity baseline, ond
ed. Sage Books, ene Colorado.

\O

HoimGren, N.H. 1997. Crataegus. In: A. Cronquist, N.H. Holmgren and P.K. Holmgren.
latetiaonataia “ie 3A, pp. 154-158.
KruscHKE, E.P. 1965 Contributions to the taxonomy of Crataegus. Milwaukee Public Mus.

Ve

Public. Boe. 3:1-27:
ove, R. and M. FRIGEN. 1978. ae hybridization ee native and naturalized
Crataegus een in western Oregon. Madrofio 25:2

NELson, A. . Crataegus wheeleri, n. sp. Bot. Gaz. 34 309.

PALMER, E.J. os Synopsis of North American Crataegus. J. Arnold Arbor.6:5—128.

Puipps, J.B. 1998. Synopsis of Crataegus Series Aptifoliae, cls yy and Brevispinae
(Rosaceae subfam. Maloideae). Ann. Missouri. Bot. Gard. 85:475—491

660 Sipa 183)
sk J.B. 1999. ae ease to the red-fruited Crataegus of western North America.
nad. J. Bot. 76:1
pars J.B. ane Gy, O "KENNON. 1998. Three new species of Crataegus (Rosaceae) from
estern North America: C. okennonii, C. okanagenensis and C. phippsii. Sida 18:169-181.
ae J.B., K.R. Rosertson, P.G. SmMiTu, R. ce 1990. A checklist of sub-fam-
ily eee (Rosaceae). Canad. J. Bot. 68:2209-2269.
SARGENT, C.S. 1889. Crataegus douglasii L an var. rvvilaris, n. var. Gard. & Forest 2:401.
SARGENT, C.S. 1890. The Silva of North America, [V:87—88, t. 176. Houghton Mifflin,
Boston, cae a New York, N
SARGENT, C.S . The Silva of Nowh America. Suppl. XHI: 37-38, t. 636. Houghton
Mifflin, ee an and New York, N
SALLABANKS, R. 1993. Fruiting plant attractiveness to avian seed dispersers: native vs. in-

DH
SS

vasive Crataegus in western Oregon. Madrofio 40:108—116.

Torrey, J. and A. Gray. 1843. Crataegus rivularis. In: A Flora of North America 1:464.,
New York, Wiley & Putnam.

Weber, W.A. and R.C. Wirtmann. 1992. Catalog of the Colorado Flora. Univ. of Colo-
rado Press, Niw

WetsH, S.L. 1982. Ys In: Utah Flora, Rosaceae. Great Basin Naturalist. 42:9—

THE SEGREGATES OF SENECIO, S.L., AND
CACALIA, S.L., IN THE FLORA OF NORTH
AMERICA NORTH OF MEXICO

THEODORE M. BARKLEY!

Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
barkley@brit.org
ABSTRACT
The traditional, broadly conceived genera Senecio, s.1., and Cacalia, s.1., will be divided
into several segregate genera in the forthcoming Flora of North America North of Mexico,
(FNANM). Provided here are a synoptical key and annotated catalog for the genera, and a
nomenclator for the specific and infraspecific epithets for Senecio, s.1., and Cacalia, s.1., that
have been commonly used in North American floristics
RESUMEN
Los géneros, tradicionalmente concebidos ampliamente, Senecio, s.1., y Cacalia, s.1., serdn
divididos en varios géneros segregados en la Flora of North America North of Mexico, (FNANM)

de préxima aparici6n. Ofrecemos aqui una clave sinéptica, un catélogo anotado de los géneros
y un indice de los epitetos especificos e infraespectficos de Senecio, s.1., y Cacalia, s.l., que

han sido usados normalmente en las floras norteamericanas

In 1978, an account of the Senecioneae in North America north of Mexico
was published in the North American Flora series (NAF) of The New York
Botanical Garden, and therein Barkley (1978) and Pippen (1978), respec-
tively, treated Senecio and Cacalia in the traditional, inclusive senses that
derive from the works of Bentham (1873a, b) and Hoffmann (1892). In so
doing, they were in agreement with the floristic botany of North American
tradition. Since the time of the NAF publication, new information and rigorous
phyletic notions of genera have combined to justify the acceptance of a greater
number of more narrowly circumscribed genera. The notions leading to these
narrower generic concepts are noted in Bremer (1994) and in several papers
that were presented at the Compositae Conference at Kew in the summer
of 1994 (Hind & Beentje 1996), particularly the paper by Barkley et al.
(1996). Preparation of the treatments of Senecio, s.1., and Cacalia s.1., for Flora
of North America North of Mexico (FNANM) has drawn attention to the mat-
ters of generic concepts, for a goal of the FNANM is to reflect current un-
derstanding as best as possible. In this paper I describe how the native and

‘Professor emeritus, Kansas State University, Manhattan, KS 66506.

Sipa 18(3): 661-672. 1999

662 Sipa 18(3)

naturalized members of Senecio, s.1., and Cacalia, s.1., will be treated generi-
cally in che FNANM. Included is a key to the genera that are being recog-
nized in the FNANM, an annotated catalog of the genera, plus a list of the
species and infraspecific names that have been commonly used in recent floristic
works, with their dispositions among the recognized genera.

Some of the genera recognized here are similar to infrageneric groups
that have been used in the past, e.g., Bentham (1873 a, b) Greenman (1901),
Pippen (1968), and Barkley (1978). ). What is new in North American floristic
work is treating them as genera, rather than as aes or sections, or as
informal groups within Senecio, s.1., or Cacalia, s.1. Morphological intergra-
dation among many species of Senecio, s.1., has been documented repeatedly
(cf. Barkley 1988 for species here meetuedl in Packera);, so far as I am aware,
however, there is no intergradation between any pair of species that are treated
here in different genera, i.e., the genera are biologically discrete.

Apart from the small and distinctive subtribe Blennospermatinae Rydb.,
the Senecioneae appear to be naturally divisable into two evolutionary lin-
eages, the Tussilagininae (Cass.) Dumort. and the Senecioninae (Cass.) Dumort.
The distinctions between these two lineages are noted in the first couplet
of the key, and they are summarized by Bremer (1994). The Tussilagininae
(“tussilaginoids”) have been called the “cacalioids” and the “tephroseroids”
in the past. Application of the name Caca/ia has been confusing, with some
consensus that it is best used for a group of Eurasian plants that are senecionoid
in their affinities. Continued uncertainty about Cacal/ia has led to its for-
mal rejection (Wagenitz 1995; Brummict 1998), and so referring to the
group as “cacalioid” is also rejected

The key to genera presented here is essentially a synopsis rather than a
practical key for identifying a plant-in-hand. No one doubts the biological
significance of the first couplet in the key, but the characters used there are
impractical for routine plant identification (Barkley et al. 1996). A key for
easy identification will need to be a “collective key” that is adeniteedly artificial
and that includes species of several genera. Such a key will be similar in
structure to the species keys for Sewecio, s.1., and Cacalia, s.1., in traditional
floristic works of the recent past.

KEY TO GENERA
1. Stigmatic areas confluent on adaxial faces of style branches; anther collars
cylindrical and cells not inflated; mostly n = 30 or polyploid/dysploid de-
rivatives. (Tussilagininae)
Shrubs with leaves concentrated distally on branches; AZ, NM... Ll. Barkleyanthus
eas rbs with principal leaves basal and on proximal half of stem.

N N

. Corollas yellow; capicula radiate or discoic

4. Capitula discoid; phyllarics notably ye ligies WY. 2. Yermo

~Capitula radiate; phyllaries green or grayish green to dirty white.

BARKLEY, Segregates of Senecio and Cacalia

5. Principal leaves with blades lanceolate to ovate, pinnately veined
and tapering to a winged petiole; boreal, WY and northward.

. Tephroseris
5. Eneye) leaves mace one rounded to reniform, plamately or
subpalm ly y contrac ted toanu nwinged petiole.
6. Flexible, oe herbs, ately exceeding 3 dm an blades of prin-
cipal leaves 1—2.5 cm across; BC. 4, Sinosenecio
6. Coarse, ae ligneous herbs to 12 dm tall; blades of princi-
pal leaves 8—16(—20) cm across; AZ (NM) 5. Roldana

3. Corollas white ee white, or ochroleucous to slightly pinkish or greenish;

capitula disc

Ts Caylee ccs elongate, racemiform clusters; Aleutian Islands.
). Parasenecio

ae Capitulescences flat-tcopped, corymbiform clusters. Southern half of
flora region.
8. Corolla lobes parted the whole lengt th of the limb, the throat indistrinct;
leaf blades deeply pinnatisect; AZ. . Psacalium
8. Corolla lobes separate less than half the length of the limb, the
hroat cylindrical or funneltorm, distinct; leaf blades subentire to

merely toothed.
9. Florets 20-40; high Appalachians (above 1400 m) NC, TN.
8. Rugelia

9. Florets 5. Widespread in southeastern quarter of the region, below
J0 m 9.

Arnoglossum
. Stigmatic areas marginal and distinc listal, adaxial faces of style branches;
anther collars swollen ie aa with basal cells inflated; n = (10) 20,

or 23, or polyploid derivatives (n = 30 in Pericallis). (Senecioninae).
). Scandent, scrambling
11. Capicula radiate; ie orange to brick-colored; apices of style branches

with distinctive, elongate central appendages; FL. ........... 10. Pseudogynoxys
1. Capitula discoid; corollas yellow; apices of style branches truncate-
penicillate; 1. Delairea
0. Plants erect and free standing.
12. Corollas white or dirty white; capitula discoid. 12. Hasteola

12. Corollas variously yellow to orange or anthocyanic, rarely white and
then capitula radiate; capitula discoid or radiate.
13. Principal leaves with blades periately veined and petioles with
clasping stipule-like | ometimes white,
or both: capitula radiate; CA 13. Pericallis
13. Principal leaves with blades not both palmately veined and with
expanded-clasping petiole bases; corollas mostly yellow, some-
times orange or ochroleucous, rarely anthocyanic; capitula dis-

coid or radiate; widespread
14. Herbs with stems arising singly or clustered from a taproot,

caudex, or rhizome and with abundant, thin, branching fibrous

roots; principal leaves in a basal cluster, cauline leaves pro-

ae reduced atsrally, margins without conspicuous callose

ticles; n = 22 or 23, or polyploid derivatives. ........00. 14. Packera

14. Herbs, subshrubs or shel of varicus habits and leaf dis-

positions, but if herbs with stems arising from a cluster of

~)

664 Stipa 18(3)

basal leaves and with cauline leaves progressively reduced
seams then the roots are fleshy and unbranched and/or the
margins have abundant callose denticles; n=(10) 20, or
polyploid derivatives. 15. Senecio (s. str.)

THE TUSSILAGINOID GENERA
1. Barkleyanthus H. Rob. & Brettell
One species; a semi-weedy, hairless shrub that occurs from Honduras northward
through Guatemala and Mexico and barely enters the FNANM range in
southern Arizona and New Mexico. Barkleyanthus salicifolins (A.B.K.) H.
Rob. & Brettell was widely known as Senecio salignus DC. in the past (Robinson
& Brettell 1974). Included in Senecio by Barkley (1978)

2. Yermo Dorn

One species, Yermo xanthocephala Dorn, is known only from barren desert
sites in central Wyoming. It was discovered and described in the past de-
cade (Dorn 1991) and its phyletic affinities are not yet well understood.
Superficially, ic rests easily in the tussilaginoid assemblage. This distinc-
tive species was given a color photograph on the cover of the magazine “Science
News” 155(1), January 2, 1999. It was not treated by Barkley (1978) or
Pippen (1978).

3. Tephroseris Rchb.

A group of some 40—50 boreal and alpine herbs, centered in Eurasia but
with at least four species in the FNANM region. The group was revised by
Cufodontis (1933), and since his time it has been treated chiefly in floristic
studies with differing species concepts (e.g., Barkley 1978; Cody 1996, Hultén
1950 & 1968; Schischkin 1968; Scoggan 1979; Welsh 1974). The taxonomy
of the group is complicated by intergradant species and unresolved nomen-
clatural problems. The species of Tephroseris superficially resemble certain
species of Senecio, s. str., but the microcharacters are clearly tussilaginoid
and the basal chromosome number is x=24, a number otherwise unknown
in the Senecionoid lineage. Included in Senecio by Barkley (1978).

4. Sinosenecio B. Nord.

A group of about 30 species of low herbs, mostly of eastern and southeast-
ern Asia, but with a single species in the New World, Sznosenecio newcombei
(Greene) J.P. Janovec & T.M. Barkley. It is apparently restricted to the Queen
Charlotte Islands, British Columbia Jlanovec & Barkley 1996). Included
in Senecio by Barkley (1978).

5. Roldana LaLlave

Fifty or more species of coarse herbs, shrubs, and small trees, most of which
have broad, palmately or subpalmately veined leaves. The group is centered
in the Trans-Mexican Volcanic Belt; it ranges through much of Mexico and

BarKLey, Segregates of Senecio and Cacalia 665

Central America. One species, Ro/dana hartwegit (Benth.) H. Rob. & Brettell,
occurs northward to southern Arizona. The genus is under revision by A.
Michele Funston. Included in Senecio by Barkley (1978).

6. Parasenecio W.W. Sm. & J. Small

A group of about 50 species, centered in eastern Asia, but one species, Parasenecio
auriculata (A. DC.) J.R. Grant, barely enters the FNANM ran ge on five islands
at the western end of the Aleutian Island chain in Alaska. Our plant has
been variously included in the past within Cacalia, Koyamacalia, Ligularia,
or Parasenecio (Grant 1993). It was not treated by either Barkley (1978) or
Pippen (1978).

7. Psacalium Cass.

About 45 species, mostly in Mexico, but one species, Psacalinm decompositum
(A. Gray) H. Rob. & Brettell, enters the FNANM region in southern Ari-
zona. The group was treated by Pippen (1968), who included our plant in
the segregate genus Odontotrichum, cf. Robinson and Brettell (1973). Included
in Cacalia by Pippen (1978).

8. Rugelia R.S. Shuttlew. ex Chapm.

A single species, Rugelia nudicaulis Shuttlew. ex Chapm. occurs in the high
mountains of central Appalachia along the Tennessee-North Carolina bor-
der. It has been treated as Senecio rugelia A. Gray in many floristic works of
the past, but was included in Cacalia by Pippen (1978).

9. Arnoglossum Raf.

Eight species of tall herbs of the southeastern quarter of the FNANM re-
gion. They form the traditional core of Cacalia, as that genus has been con-
ceived in North American floristics. A new species was recently described
by Anderson (1998). Included in Cacalia by Pippen (1978).

THE SENECIONOID GENERA

10. Pseudogynoxys (Greenm.) Cabrera

A dozen or so species; scandent, scrambling vines with showy, orange or
brick-colored corollas. They are native to the Caribbean region and south-
ward as far as eastern Brazil. They are cultivated as ornamentals, and one
species, Psendogynoxys chenopodtoides (Kunth) Cabrera, escapes from cultiva-
tion and persists in southern Florida and perhaps elsewhere along the Gulf
Coast. It has been called Senecio confusus Britten in floristic works and the
horticultural literature. The group was catalogued and a key was presented
by Robinson and Cuatrecasas (1977), but the relationships among the spe-
cies remain poorly understood and a revision is needed. Included in Senecio

by Barkley (1978).

666 Sipa 18(3)
L1. Delairea Lem.

One species; a scandent vine, Delairea odorata Lemaire, 1s native to southern
Africa but is now established as an aggressive weed in coastal California.
This plant has been known as Senecio mikanioides Otto, and was included in
Senecio by Barkley 1978.

12. Hasteola Raf.

Two species; one is of the east-central U.S and the other was recently de-
scribed from Florida. These plants have eradiate capitula and white corol-
las, which places them in the traditional concept of Cacalia, but they have
the microcharacters and cytology of the Senecionoid lineage, cf. Anderson
(1994). Included in Cacalia by Pippen (1978).

13. Pericallis D. Don

About a dozen species; native to the Canary Islands, Madeira, and the Azores,
and some species are ancestral to the horticultural complex called “the florists’
cineraria.” A cultivar from the complex escapes and persists in favored sites
in coastal California. Under current taxonomic concepts our plant is best
treated as Pericallis hybrida B. Nord., but it has been widely known in the
past as Senecio cruentus DC. Pericallis is alone among the Senecionoid genera
in having n = 30,a number that is typical of the Tussilaginoid lineage. In
all other characters, however, it fits comfortably among the Senecionoids
(Nordenstam 1977, 1978). Included in Senecio by Barkley (1978).

14. Packera A. Love & D. Live

About 60 species; centered in the western temperate half of North America
but extending into southern Mexico and into the Arctic. Two arctic-alpine
species extend into northern Siberia and central Asia. . The group has been
known as the “aureoid complex” of Senecio, s.1., and although its members
are superficially similar to many species of Seveczo, it forms a distinct lin-
eage. Intergradation among Packera species is well known (Barkley 1988),

but there is no morphological intergradation or putative hybridization be-
tween any Packera and any species of Senecio, s.str., or segregate of Senecio. In
addition to the characters used in the key, Packera is further distinguished
from Senecio by having pollen grains of a helianthoid ultrustructural type
rather than of a senecionoid type (Bain & Walker 1995). An ITS-based phylogeny
has been estimated for Packera by Bain and Jansen (1995). Included in Senecio

by Barkley (1978).

15. Senecio L.

Senecio, in the broadest sense, is a nearly world wide genus of some 3,000
species, and even after the exclusion of the currently accepted segregates, it

still encompasses some | ,300 species. Clearly, Senecio, s.str., is a “mixed bag”

BaRKLEY, Segregates of Senecio and Cacalia 667

that is defined largely as what remains after the rather precisely circum-
scribed segregate genera are removed. It includes numerous species of un-
known affinity, and presumably the concept of “Senecio” will change as more
is learned about the relationships of the currently included species. About
50 species of Senecio, s.str., occur in the FNANM region; some are native,
some are introduced, and a few are notable weeds.

SENECIO AND CACALIA NAMES OF TRADITIONAL USE IN NORTH AMERICAN
FLORISTICS AND THEIR DISPOSITION IN THE FNANM

Epithets with no alternative name remain as listed
SENECIO
actinella Greene
amplectens A. Gray

var. amplectens

var. holmii (Greene) H.D. Harr.
ampullaceus Hook.
anonymus A.W. Wood = Packera anonyma (A.W. Wood) W.A. Weber & A. Love
antennartifolius Britton = Packera antennariifolia (Britton) W.A. Weber & A. Love
aphanactis Greene
arizonicus Greene
aronicoides DC.
astephanus Greene
atratus Gree
inane (Led eb.) Fedtsch. = Tephroseris urea (Ledeb.) om subsp. atropurpurea

var. frigidus = T. a. subsp. frigida oo ‘A. Lave &

var. tomentosus (Kjellm.) Awan - oo ala (A.E. Bei Holub
aureus L. = Packera aurea (L.) A
bernardinus Greene = Packera ae i ae W.A. Weber & A. Live
bigelovii A. Gray

blochmaniae Greene
bolanderi A A. Gray = Packera bolandert (A. Gray) W.A. Weber & A. Love
var. bolanderi
var. harfordii (Greenm.) T.M. Barkley = P. db. var. harfordii (Greenm.) D.K. Trock & T.M.

Barkley
brewert Burtt = Packera breweri (Burtt Davy) W.A. Weber & A. Love
californicus DC
cannabifolins Less.
canus Hook. = Packera cana (Hook.) W.A. Weber & A. Love
cardamine Greene = Packera cardamine (Greene) W.A. Weber & A. Léve
castoreus 8.L. Welsh (a Packera?)
clarkianus A. Gray
clevelandii Greene = Packera clevelandii (Greene) W.A. Weber & A. Love
confusus Britten = Pseudogynoxys chenopodioides (Kunth) Cabrera
congestus (R.Br.) DC
conterminus Greenm. = Packera contermina (Greenm.) J.F. Bain

668 SIDA 18(3)

crassulus A. Gray

crocatus Rydb. = Packera crocata oe) W.A. Weber & A. Love

cruentus DC. = Pericallis hybrida B. Nor

cymbalaria Pursh = Packera cymbalaria (Pursl 1) W.A. Weber & A. Live

cymbalarivides Buek = Packera buekti D.K. Trock & T.M. Barkley

Bene Greene = Packera cynthotides eee W.A. Weber & oa Love
debilis Nutt. = Packera debilis (Nutt.) W.A. Weber & A. L

Lint sreene = Packera Linoihiby /la (Greene) WA. Weber & A. Love

limorphophylla

var. intermedius T. Barkley = P. d. var. intermedia (T.M. Barkley) D.K. Trock & T.M. Barkley
var. paysonii T. Barkley = P. d. var. paysonii (T.M. Barkley) D.K. Trock & T.M. Barkley

douglasii DC. = Senecio flaccidus Less.
var. douglasii = S. f. var. — — .) B.L. Turner & T.M. Barkley
var. longilobus (Benth.) L.D. Ben = S. f Less. var. flaccidus
var. monoensts (Greene) Jeps. = $ 2 var. monoensis (Greene) B.L. Turner & T.M. Barkley

elegans L

elmer? Piper

eremophilus Richardson

var. ere mophi

var. kingii (Ry. db.) Greenm
var. macdougallii (A. Helle r) Cronquist

ertterae T.M. Barkley

ica Torr. & A. Gray = Packera eurycephala (Torr. & A. Gray) W.A. Weber & A. Live

eurycephala
var. pe isroset (J.T. Howell) T.M. Barkley = P. ewrycephala. var. lewisroset J.T. Howell)
J.-E Bain

fendleri A. Gray = Packera fendleri (A. Gray) W.A. Weber & A. Love

flaccidus Less.
var. flaccidus
var. douglasit (DC.) B.L. Turner & T.M. Barkley
var. monoensis (Greene) B.L. Turner & T.M. Barkley

flettii Wiegand = Packera flettii (Wiegand) W.A. Weber & A. Live

foetidus Howell = Senecio hydrophiloides Rydb.

foetidus var. hydr alias (Rydb.) T.M. Barkley ex Cronquist = eae eas ophiloides Rydb.

franciscanus Greene ackera franciscana (Greene) W.A. Weber

fremonti Torr. & A. ah
var. fremontit
var. blitoides (Greene) Cronquist
var. occidentalis A. Gray
var. /nexpectans Cronquist

frigidus Less. = Tephroseris atropurpurea subsp. fridiga emit A. Live & D. Love

fuscatus Hayek = Tephroseris lindstroemii A. Live & D. Live

| T.M. Barkley & R.M. Beauch. = Packeri ganderi (1.M. Barkley & R.M. Beauch.)

. Weber & A. Lévy
aa Poir. kera clabella (Poir.) C. Jeffrey
ereenet A, Gray = Packera greenei (A. Gray) WA, Weber & A. Live

hartianus A. ae = Packera hartiana ae Heller) W.A. Weber & A. Live
artweeti Benth. = Roldana hartwegii (Benth.) H. Rob. & Brettell
ae rius Greene = Packera hesperia (Greene) W.A. Weber & A. Live
huachucanus A. Gray = Senecio multidentatus var. huachucanus (A. Gray) T.M. Barkley

669

BarKLey, Segregates of Senecio and Cacalia

hydrophiloides a

Aydrophilus N
hyperborealis an = Packera hyperborealis (Greenm. ) A. Love & D. Live
imparipinnatus Klatt = Packera tampicana ( . Jeffrey

indecorus Greene = Packera indecora (Greene) A Love & D. Love
integerrimus Nutt.
var. sntegerrimits
var. exaltatus (Nutt.) Cronquist
var. major (A. Gray) Cronquist
var. ochrolencous (A. Gray) Cronquist
var. scribneri (Rydb.) T.M. Barkle
oe Sea Greene = Packera ionophylla (Greene) W.A. Weber & A. Live
jacobaee
ie A E. Porsild = Tephroseris kjellmanii (A.E. Porsild) Holub
layneae Greene = Packera layneae (Greene) W.A. Weber & A. Live
lemmonii A. Gray
lindstroemti A.E. Porsild = Tephroserts lindstroemii (A.E. Porsild) A. Live & D. Live
/ugens Richardso
onil c
Macounit (Gee = Packera macounii (Greene) W.A. Weber & A. Love
malmstenti §.F. Blake ex Tidestr. (a Packera ?)
megacephalus Nutt.
mikanioides Otto ex Walpers = Delairea odorata Lem.
millefolium T.& G. = Packera millefolia (Torr. & A. Gray) W.A. Weber & A. Live
millelobatus was b. = Packera millelobata (Torr. & A. Gray) W.A. Weber & A. Love
mohavensis A. Gra
ee (Calder & R.L. Taylor) G.W. Douglas & G. Ruyle-Douglas =
alder & R.L. aes FE. Bain
Yin: Greenm. = Senecio a Gee var. multicapitatus (Greenm.) S.L. Welsh
ray) T.M. Bark y

L pis
3 LOPOSUECHES L

multidentatus var. ok aia (A.G
baie Torr. & A. Gray ex A. Gray = Packera multilobata (Torr. & A. Gray ex A. Gray)
Weber & A. Love
MUSIBIENS IS oy L. Welsh (a Packera
neomexicanus A. Gray = Packera neomexicana (A. Gray) W.A. Weber & A. Live
var. neomexicanus = P. n. var. neomexicana

var. prods (Greene) T.M. Barkley = P. n. var. metcalfei (Greene) D.K. Trock & T.M.

var. mee (Greene) T.M. Barkley = P. ». var. mutabilis (Greene) W.A. Weber &
A. Lov

var. cee (Greene) T.M. Barkley = P. n. var. towmey? (Greene) T.M. Barkley & D.K.

Troc
neowebsteri S.E. Blake
newcomber Greene = Sinosenecio newcombei (Greene) J.P. Janovec & T.M. Barkley
obovatus Muhl. ex Willd. = Packera obovata (Muhl.ex Willd.) W.A. Weber & A. Live
ogotorukensis Packer = Packera ogotorukensis (Packer) W.A. Weber & A. Love
parryi A. Gray
pattersonensis Hoover
pauciflorus Pursh = Packera pauciflora (Pursh) W.A. Weber & A. Love
pauperculus Michx. = Packera paupercula (Michx.) W.A. Weber & A. Live
plattensis Nutt. = Packera plattensis (Nutt.) W.A. Weber & A. Live

670 Sipa 18)

porteri Greene = Packera porteri (Greene) C. Jeffrey
psendaureus Rydb. = Packera sasaki — ) W.A. Weber & A. Love

var. psendaurens = P. p. var. pseue daur

var. flavulus (Greene) tes = =p, p. var. flavula (Greene) W.A. Weber & A. Love

var. ee a & Bush) T.M. Barkley = P. p. var. semicordataus (Mack. & Bush)
TM y & D.K. Troc ck

pseudo-arnica a
pudicus Greene
quaerens Greene = Packera quaerens (Greene) W.A. Weber & A. Live
quercetorum Greene = Packera quercetorum (Greene) C. Jeffrey
raptfolius Nutt
seas Less. = Packera cymbalaria (Pursh) W.A. Weber & A. Love

riddelli Torr. & A. Gray
vobbinsi Oakes ex Rusby = Packera schweintiziana (Nutt.) W.A. Weber & A. Live
rugelia A. Gray = = Rugelia nudicaulis Shutlew. ex Chapm.
sacramentanus Wooton & Stand.
salignus DC. = Barkleyanthus salicifolius (Runth) H. Rob. & Brettell
sanguisorboides Rydb. = Packera sanguisorboides (Rydb.) W.A. Weber & A. Live
schweinitzianus Nutt. = Packera schweinitzianus (Nutt.) W.A. Weber & A. Love
scorzonella Greene

Hook

SOVVA

var.

var. nalts (Greene) A. Nelson
sheldonensis A.E. Porsilc
smallii Britt. = Packera anonyma (A.W. Wood) W.A. Weber & A. Léve
soldanella A. Gray
sSpartioides Torr. & A. Gray

var. spartioides

var. r. multicapitatus (Greenm.) S.L. Welsh
spellenbergii T.M. Barkley = Packera spellenbergti (T.M. Barkley) C. Jeffrey
sphaerocephalus Greene
streptanthifolius Greene = Packera streptanthifolia (Greene) W.A. Weber & A. Live
subnudus DC. = Packera buekii D.K. Trock & T.M. Barkley

e

tampicanus DC. = Packera ene (DC.) C. Jeffrey

taraxacoides (A. Gray) Gre

tomentosus Michx. = Pockets tomentosa (Michx.) C. Jettrey

triangularis Hook.

tridenticulatus Rydb. = Packera tridenticulata (Rydb.) W.A. Weber & A. Live
VISCOSHS 7

vilearts L.

warnocki? Shinne

wer oe A. Gr = Packera werneriifolia (A. Gray) W.A. Weber & A. Live
wootont Gre

yukonensts A. E. "Porsild = Tephroserts yuRonensis (A.E. Porsild) Holub

CACALIA

auritulata D arasenecio auriculata R. Grant
decompostta A. Gray = = Psacalinm decompositum (A. Gray) H. Rob. & Brettell

atriplicifolia L. = Arnoglosum atriplicifoliium (L.) H. Rob.
= P. Cs

BARKLEY, Segregates of Senecio and Cacalia 671

diversifolia Torr. & A. Gray = Arnoglossum diversifolium (Torr. & Gray) H. Rob.

floridana A. = Arnoglossum floridanum (A. Gray) H. Rob.

ei scrres an Bip. = Arnoglossum cies me (Hook.) H. Rob.

ovata Walt. = = Arnoglossum ovatum (Walte Rob.

ee (Raf.) Shinners = ae pam Raf.
ugelia (Shuttlew. ex Chapm.) T. Barkley & Cronguist = Rugelia nudicaulis Shuttlew. ex

Chapm.
suaveolens L. = Hasteola suaveolens (L.) Pojark.
sulcata Fernald = Arnoglossum sulcatum (Fernald) H. Rob.

ACKNOWLEDGMENTS

Conversations on the matters of generic delimitations in the Senecioneae
were had with many people over the years, including Loran Anderson, John
Bain, Luc Brouillet, Bonnie Clark, Craig Freeman, Michele Funston, Hugh
Itis, Charles Jeffrey, Robert Kowal, Alison Mahoney, Guy Nesom, Bertil
Nordenstam, John Packer, Harold Robinson, John Strother, Debra Trock,
Billie Turner, Jose Luis Villasefior, William Weber, and Alan Whittemore.
It is my pleasure to acknowledge all of them. John Strother and Robert Kowal
were also of great assistance in editorial and nomenclatural matters. It is
also a pleasure to tip the hat to the memory of the late Arthur Cronquist,
who so gladly shared with me his great knowledge of the North American
Compositae.

REFERENCES

ANDERSON, L. 1994. * revision of Hasteola (Asteraceae) in the New World. Syst. Bot. 19:211—219.

ossum album (Asteraceae): New species from northern Florida.

Sida rane ee
Bain. J.F. and R.K. ve 1995. A phy eae analysis sa aureoid Senecio (Asteraceae)
—219,

complex ee on ITS sequence data. Pl. Syst. Evol. 195:2
WALKER 1995. A comparison of the mie wall ultrastructure of
aureod and non-aueroid Senecio species (Asteraceae) in North America. Pl. Syst. Evol.

LU.

BENTHAM, G. 1873a. Notes on the classification, ie and geographical distribution of
the Compositae. - ee Soc., Bot. 13:335-57

3b. Compan ce In: G. eee & J.D. Hooker, Genera Plantarum,

vol.2, part 1. o. & Co. and Williams & Norgate, London.

Bac TM. 1978. Senecio. N. Amer. FI. IL. 10:50-139.

O88. V: ‘ : g } | _ ENT.
perspective. Bot. Rev. 54:82—106.

BarKLEY, T.M., B.L. Clark, and A.M. Funston. 1996. The segregate genera of Senecio, s.1.,
and Cacalia, s.1. (Asteraceae: Senecioneae) in Mexico and Central America. In: D.J.N
Hind and H.J. Beentje, eds. Compositae: Systematics. Proceedings of the International
Compositae Conference, Kew 1994. 1:613—620. Royal Botanic Gardens, Kew, U.K.

Bremer, K. 1994. Asteraceae: Cladistics and classification. Timber Press, Portland, OR.

h America: A geohistorical

Brummit, R.K. 1998. In: Nicolson, D.H., ed. Proposals to conserve or Reject. Report on
the Committee for Spermatophyta. Taxon 47:444.

672 Sipa 18(3)

Copy, W.J. 1996. Flora of the Yukon Territory. NRC Research Press. Ottawa, Canada.

Curopontis, G. 1933. Kritische Revision von Senecio sectio Tephroseris. Feddes Repert. Spec.
Nov. mee 70: 1-266.

Dorn. L991. Yermo xanthocephalus eas Senecioneae): A new genus and species
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GRANT, J.R. 1993. A new cone et os ee nn single A | f Para: 10 (A
Senecioneae). Novon 3:154—155

GREENMAN, J.M. L901. Monographie der nord- und centralamerikanischen Arten der Gattung
Senecio. 1. Allgemeines und Morphologie. Leipzig. (Reprinted in Bot. Jahrb. Syst. 32:1—
33. 1902.)

Hinp, D.J.N. and H.J. pene (eds.). 1996. Compositae: Systematics. Proceedings of the
International Compositae Conference , Kew, 1994. Royal Botanic Gardens, Kew, U.
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45, Nr. 1, pp. 1601-1623. C.W.K. Gleerup, Lund, Sweden

a 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press,
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Janovec, J.P. and T.M. BarKLEy. 1996. Sinosenecio newcomber (Asteraceae: Senecioneae): A
new combination for a North American plant in an Asiatic genus. Novon 6:265—267.

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1978. Taxonomic notes on the tribe Senecioneae (Compositae). Opera
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PIPPEN, 1968. aes ‘Cacalioid” genera allied to Sevecio (Compositae).
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1978. a N. Amer. FI. IL. 10:15 1-159.

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oy

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A NEW VARIETY OF PHYSARIA NEWBERRYI
(BRASSICACEAE) FROM NEW MEXICO

ROBERT C. SIVINSKI
New Mexico Forestry Division
.O, Box 1948
Santa Fe, NM 87504, U.S.A.
bsivinski@ state.nm. us
ANG
Herbarium
Department of Biology
University of New Mexico
Albuquerque, NM 87131-1091, U.S.A.

ABSTRACT
The new variety Physaria newberryi var. yesicola is described and illustrated. It is readily

distinguished within P. newberryi by its very long styles. Variety yesécola occurs upon the
gypseous Yeso Formation in the Sierra Lucero of west-central New Mexico.

RESUMEN
Se describe y se ilustra una nueva variedad Physaria newberryi var. yesicola. Se distingue

facilmente de P. newberryi por sus estilos muy largos. La variedad yesicola aparece en suelos
gipsicolas de la Formacién Yeso en la Sierra Lucero del centro-oeste de Nuevo México.

Physaria newberryi A. Gray var. yesicola Sivinski, var. nov. (Fig. 1). Tyee:
U.S.A. NEW MEXICO. Valencia i. NE side of Sierra Lucero ca 37 km W of Los
Lunas, T6N R3W Section 2 NW'/4 NW!/4, 34°46'54"N_ 107°7'48"W, elev. 1800

4 May 1998, R. Sivinski & C. eke 35 (HOLOTYPE: UNM; isotypes: ARIZ,
BRY, CAS, GH, MO, NY, US).

A Physaria newberryi A. Gray var. racemosa Rollins stylis filiformibus (S—9 mm longis) et
racemis fructiferis brevibus (2.5—5 cm longis) differt.

Long-lived caespitose perennial with diffusely branching caudex forming mounded
clumps 10-30 cm in diameter; caudex branches thickly clothed with marcescent
leaf bases and terminated by clusters of ascending to erect leaves; stems and
leaves densely covered with overlapping stellate-discoid trichomes, trichome
rays confluent for at least 1/2 their length and often to near their apex; basal
leaves narrowly oblanceolate to broadly spatulate, 3-8 cm long (including
petiole), acute to obtuse, margins entire or with a few broad teeth, the winged
petiole less than to 2 times as long as the expanded blade; cauline leaves
few, sessile, 1-1.5 cm long, linear-oblanceolate; stems numerous and aris-
ing from the axils of basal leaves, ascending to erect; mature racemes 2.5—5

SipA 18(3): 673—677. 1999

674 Sipa 18(3)

» AR
a Ve oe

Fic. 1. Physaria newherryi var. yesicola Sivinski var. nov., (A) growth form (one-half of cae-
spitose se (B) detail of plant and latitudinal cross-section of silicle from Sivinski &
McDonald 433
cm long; faalehap pedicels straight, ascending or divergent, 6—11 mm long;
sepals greenish yellow, pubescent, lanceolate and somewhat cucullate, 6—7
mm long and 1.0-1.5 mm wide; petals yellow, oblanceolate to spatulate,
7-8 mm long and 1.5—2.0 mm wide; sé/icles papery, pubescent, becoming
purplish at maturity, deeply cleft at the apex with a V-shaped sinus and with
little or no basal cleft, fruit cross-section X-shaped with concave valve sides and
dorsal surface, valve margins and sinus crests sharply keeled, each valve 6—9 mm
long and 4-7 mm wide; repla 2.5—3.5 mm long, narrow to nearly closed, acute
at the apex; fruiting styles thread-like, 5-9 mm long, surpassing the apical cleft of
the silicle; ovules 2 per valve; seeds ovate, slightly compressed.

Additional specimens examined: NEW MEXICO: Cibola Co.: canyon on the west side
of Sierra Lucero, ca 47 km S of Laguna biccases) TSN RSW Section 36 NE'/s, 34°37'13"N

107°18'5S"W, elev, 2040 m, 14 May 1998, R.C. Sivinski 4368 (ASC, COLO, ISTC, NMC,

RENO, RM, UNM, UT). Valencia Co.: icc Pueblo Reservation, NE side of Mesa Lucero,
34°50'14"N, 107°6'43"W, elev. 1740 m, 24 Sep 1997, R. Sivinski, T. Lowrey & B. Miller
4160 (UNM).

Distribution and Habitat.—Variety yesicola is presently known only from
the Sierra Lucero Range (including Mesa Lucero) of Cibola and Valencia

SIVINSKI, A new variety of Physaria newberryi 675

counties in west-central New Mexico (Fig. 2). This population is the ex-
treme southeastern range limit for the species. It occurs on sandy gypsum
and other silty strata of the Permian age Yeso Formation. The Yeso Forma-
tion is 200 to 300 meters thick in the Sierra Lucero. It is comprised of a
soft, silty sandstone interbedded with gypsum, limestone, shale and silt-
stone strata of various thicknesses (Weber & Kottlowski 1959). Variety yesicola
occurs on silty sand substrates that contain obvious quantities of gypsum.
It is also locally abundant on adjacent siltstone and silty limestone strata
which may be mildly gypseous, but have not been analyzed.

The habitat is nearly barren badlands and canyon sides of various slopes
and exposures between the elevations of 1700 and 2100 m. Juniperus monosperma
(Engelm.) Sarg. trees are scattered across this formation and the s
and herbaceous vegetation varies with the geologic strata. Common associ-
ates on sandy gypsum are T7quilia hispidissima (Torr.) A. Richardson, Se/inocarpus
lanceolatus Wooton, Calylophus hartwegii subsp. filifolius (Eastw.) Towner &
Raven, Cryptantha fulvocanescens 8. Wats., Artemisia bigelovii A. Gray, Tetradymia
fulifolia Greene, Lycium pallidum Miers, Sporobolus nealleyi Vasey and Stipa
comata Trin. & Rupr. The most frequent gypsum habitat associate is an
undescribed Phacelia that is in preparation for publication by Tim Lowrey
and Paul Knight (UNM) with Dwane Atwood (BRY). Associates on silty
limestones or siltstones are more variable, but often consist of Ephedra torreyana
S. Wats., Rhus trilobata Nutt., Eriogonum corymbosum Benth., Nolina greenet
S. Wats., Aristida purpurea vat. fendleriana (Steud.) Vasey and Schizachyrinm
neomexicanum Nash.

Taxonomic Relationships. —Physaria newberryi is broadly, but sporadically,
distributed through northwestern New Mexico, northern Arizona and southern
Utah (Rollins 1993), It is closely related to Physaria acutifolia Rydberg (Montana
to New Mexico and Utah) and Physaria chambersii Rollins of the Great Ba-
sin (Mulligan 1967). These three species can usually be distinguished with
the following key:

1. Dorsal margins of erie vanes acu angled (nearly winged); uae ape

trick r

acute valves have shed); trichome
for 1/2 or more their length P. newberryi

—

. Dorsal margins of silicle valves obtusely angled or broadly rounded; replum
apex obtuse or rounded; trichome rays confluent or free

—~
~~)
—

2. Silicles apically cleft with a deep, narrow or V-shaped sinus and with a
shallow or no basal sinus at the point of pedicel attachment, valves often
obtusely angled; trichome rays usually free (confluent in var. membranacea)

P. chambersii

in)

. Silicles narrowly cleft at both the apex and base, valves broadly rounded;
trichome rays free or confluent for nearly 1/2 their length . acutifolia

The taxonomic boundaries among these species are notably blurred by
intergrading polyploid races, especially in central and southern Utah (Mulligan

676 Stipa 18(3)

1967; Welsh et al. 1993). The distinctively angled silicle valves of Physaria
newberryi make it the most well-marked and distinguishable species of this
group (Fig. 1).

KEY TO THE VARIETIES OF PHYSARIA NEWBERRYI

1. Styles filiform, 5-9 mm long, surpassing the crest of the apical silicle sinus;
2

repla 2.5—3.5 mm long; racemes 2—5 cm long var. yesicola
1. Seyles scout, <4 mm long, usually shorter than the silicle sinus; replum and

racemes lengths not combined as abov (2)
2. Repla 4-10 mm long; racemes erie 2—5 cm long (rarely up to 10 cm)
var. newberryi

2. Repla 2—3.5 mm long; racemes 5—10 cm long var. racemosa

Variety yeszcola and var. racemosa both have short os and the latter has
also been collected from gypseous substrate (Gierish 4214, ARIZ, BYU).
Variety racemosa is a minor peripheral variant in northwestern Arizona and
southwestern Utah that possesses a unique combination of replum and raceme
lengths, but no single distinguishing morphological characteristic. Welsh
(1993) considers racemosa an insignificant phase that grades into var. newberry7.
In contrast, var. yesico/a is an isolated disjunct with long, filiform styles that
are unique within this species.

The combination of confluent trichome rays and V-shaped apical silicle
sinus in var. yesicola is also similar to P. chambersii var membranacea Rollins
(syn. P. lepidota Rollins) of south-central Utah. Yet the long slender styles
and short repla of var. yesicola are nearly as unusual for P. chambersii as they
are for P. newberryi. In this case, I place the utmost taxonomic importance
on silicle shape. The valve margins of var. yesicola are sharply keeled from
the silicle base to the crest of the apical sinus and the valve surfaces are con-
cave and less inflated than P. chambersii. These silicle features are character-
istic of P. newberryi and clearly place yesicola close to that species. The long
styles of P. newberry? var. yesicola are a conspicuous departure from the usual
circumscription of this species and further obscures the taxonomic bound-
aries between it and other related taxa. Additional study of this species group
is needed and may find justification for reducing some species to infraspecific
status within P. newberryi or else elevating var. yesicola to species level.

At present, var. yesicola is known only from the Sierra Lucero and appears
to be geographically isolated from other Physaria taxa (Fig. 2). It is locally
abundant and morphologically consistent in this 50 km range of low mountains.
This unique plant is another addition to a growing list of taxa endemic to
the gypsum formations of New Mexico.

Etymology.—This new variety dwells upon the Yeso Formation in the Sierra
Lucero, hence the name yesico/a. Yeso is the Spanish word for ‘gypsum’ which
is a fitting name for this geologic feature.

SIVINSKI, A new variety of Physaria newberryi 677

ye eo 0 50km
| t——
SAN JUAN |
|—-—-—-; |
: os 109°
McKINLEY : |
7 |
a gor eee (as eae ols
| . pa ae {| NEW MEXICO
ac CIBOLA (ne = eae
le | |
on
ee ae |

Fic. 2. Distribution of ae newberryi var. newberryi: @, P. newherryi var. yesicola: ©, and P.

acutifolia: * in New Mexi
ACKNOWLEDGMENTS

The field surveys that initially located this new taxon were funded by
grants from the U.S. Fish and Wildlife Service and the New Mexico State
Land Office. I thank the curatorial staffs at ARIZ, ASC, BRY, NMC, RM,
and UNM for making their collections available to me. John Strother and
Alan Smith kindly reviewed the Latin diagnosis. Tim Lowrey and Charlie
McDonald provided valuable field assistance and companionship.

REFERENCES

MULLIGAN, 1967. eee of ete acutifolia, P. chambersit, and P. newherryi
ae aes Can: ad. J. Bor. 45:1887—

ROLiins, R.C. 1993. The eee of peas North America. Standford Univ. Press,
Standford, CA.

Weser, R.H. and KE. Korriowsk1. 1959. Gypsum resources of New Mexico. Bulletin 68,
NM Institute of Mining and Technology, State Bureau of Mines and Mineral Resources,
Socorro.

WeELsH, S.L, N.D. Arwoon, S. Goopricn, and L.C Hicains. 1993. A Utah flora. 2nd ed.
ee Young Univ. Print Services, Provo, UT.

678 Sipa L8(3)
BOOK NOTICE

Lawrence R. HEANey and JACINTO C. REGALADO, JR. 1998. Vanishing Treasures
of the Philippine Rain Forest. (ISBN 0-914868-19-5, pbk), The Field
Museum, Roosevelt Road at Lake Shore Drive, Chicago, [linois 60605-

2496. $24 88 Pp.

This book “was published by The Field Museum, Chicago, on the occasion of “Vanish-
ing Treasures of the Philippine Rain Forest” and “Voyage a Nation: The Philippines” -two
exhibitions celebrating the biological diversity and culture of the Philippines.”

As a semi-popular work, the book provides a wonderful introduction to the diversity,
origins and prospects for survival of the highly endemic biota of the Philippines. Through
highly illustrative examples, and fantastic photographs, the reader is led through some of
the most striking elements of the remaining few extant forests. Most importantly, the bal-
ance of botanical and zoological examples of endangered taxa, and the economic conse-
quences of their habitats’ destruction, are illustrated in a practical way previously not seen
in popular books. Every reader will be impressed with the extent of forest destruction, the
catastrophic effects of deforestation on rural human populations, and mass extinction of
species that were locally common in the past. While geographic and species coverages are
slanted toward the Visayan and Mindanao regions of the country, examples are provided

from all major faunistic areas in a well-balanced manner, complete with historical and present-
day statistics. I have seen no better “coffee table” book that clearly and succinctly explains
the horrific consequences of forest destruction, endangerment of species, and degrac ation
of the environment. The case in the Philippines as the hottest of the “hot spots” is made
ina striking manner. While I disagree that the Philipy dines is “megadiverse,” (sensu Co-
lombia or Brasil), it 1s certainly “mega-enc

oo

emic” and the state of its forest cover is akin
only to the that of Madagascar or Haiti. After serious reading of this book, I can nee
imagine anyone who ease not ss ey to ae research in basic systematics, conser-

vation biology and ;

sal
jon

ment rograms in the country. It is a book that should
be on the bookshelf of every field ee conservationist and environmental enthusi-
ast.—John J. Pipoly HI.

Sipa 18(3): 678. 1999

SYNOPSIS OF NEW WORLD COMMICARPUS
(NYCTAGINACEAE)

NEIL A. HARRIMAN
Biology Department
University of Wisconsin-Oshkosh
Oshkosh, WI 54901, U.S.A.

harviman@ mwosh.edu

ABSTRACT

Two new species of Commicarpus from México (C. coctoris and C. praetermissus) are described.
A key to the five New World species is presented.

RESUMEN

Se describen dos nuevas especies de C irae ee taginaceae) de México. Se ofrece
una clave para las cinco especies del Nuevo Mund

Commicarpus Standley, predominantly an African genus of 25—30 species
(Meikle, 1978), was segregated from Boerhavia L. by Standley (1909), along
with Anulocaulis Standley (1909) and Cyphomeris Standley (1911).

Fosberg (1978) reduced Standley’s genera to subgenera under Boerhavia,
though the combinations were never formally made and therefore have no
nomenclatural standing (ICBN Art. 33.2). In modern floristic treatments,
opinion is divided as to whether the segregate genera (or at least Commicarpus)
deserve recognition. Commicarpus is recognized by, e.g., Breedlove 1986; Gonzalez
Elizondo et al. 1991; Liogier 1985; Brako and Zarucchi 1993. However,
Acevedo-Rodriguez (1996) reduces Commicarpus to Boerhavia, as does Bogle
(1974); Kellogg (in Howard 1988) likewise treats Boerhavia in the broader
sense, to include Commicarpus, but acknowledges that cladistically there is
no reason to choose one placement over the other—“We therefore include
them in the same genus for convenience.” I choose to recognize Commicarpus,
but I have chosen specific epithets for the two new species which will not
form homonyms if they are someday transferred to Boerhavia.

tudy of some general collections from southwestern México, one com-
municated by courtesy of Paul A. Fryxell, the other long overlooked in herbaria,
has revealed two novelties.

Commicarpus coctoris N.A. Harriman, sp. nov. (Fig. 1). Type: MEXICO.
AxaCA: Municipio San Agustin Atenango, 14 km al SO de San Marcos Arteaga,
carretera Huajuapan-Juxtlahuaca; laderas con matorral tropical caducifolia con unos
encinos, Brahea, Otatea, Bursera, etc., suelo calizo, somero, alt. 1770 m, hierba pegajosa

con ramas principales hasta 50 cm de largo, extendidas sobre el suelo, iealices guindas;

Stipa 18(3): 679-684. 1999

680 Stipa 18(3)

UNIVERSITY OF

; Gal
HERBARIUM r e pou :

HERBARIO—WORTORIO DEL COLEGIO |
PLA

10 efem. 2 ae,

Fic. 1. Holotype of Commicarpus coctoris N.A, Harriman.

HarRIMAN, New World Commicarpus 681

en borde del matorral, escasa, 23 Nov 1989 (flowers and fruits), Koch & Fryxell 89220
(HOLOTYPE: TEX!; isotypes: CHAPA, OSH! and others to be distributed from TEX
and CHAPA)

Haec species ob pilos glandulosos et perianthia rubra | 2-16 mm longa et lata inter omnes

species americanas peculiaris est.

Perennial herb, stems pink, to 50 cm long, sprawling over the ground,
viscid, densely beset with multicellular, spreading, gland-tipped hairs, these
mostly less than 0.5 mm long but some to 1 mm long. Leaves opposite,
entire, isophyllous, blades 30-55 mm x 15-40 mm, ovate, basally cuneate
to rounded, apically acute and cuspidate, without evident internal crystals,
marginally glandular ciliate, adaxially glabrous, abaxially glandular hairy
on the main veins only, on glandular-hairy petioles 8-12 mm long. Flow-
ering branches hairy like the main stems, terminal and from the upper ax-
ils, with pedunculate, 4—6 flowered, simple umbels. Flowers on glandular-
hairy pedicels 8-18 mm long, the portion near base of anthocarp without
stipitate glands. Perianth funnelform, cherry red, 12—16 mm high and wide,
glandular-hairy externally, glabrous internally; stamens 4, filaments gla-
brous, exserted; style glabrous, exserted well beyond the stamens, the stigma
capitate. Fruits thickly clavate, glabrous, striate but not ribbed, 6-7 mm
long, the pustular glands irregularly arrayed in the upper 1/2—2/3 of the
fruit (anthocarp).

Flowering and fruiting.—November, so far as known.

Distribution and habitat.—Known solely from the type collection, in tropical
deciduous forest. Possible paratype, the information communicated by Richard
Spellenberg, the specimen not seen by me: México. Oaxaca. Laguna Encantada,
+ 3 kmal N de Justlahuaca, colinas yesosas con vegetacion arbustiva dispersa
de Agave, Dasylirion, a alt. 1600 m, 8. Zamudio & G. Ocampo 10951,
22 Nov 1998 CEB, NMC).

Ety mology. —The epithet honors Stephen D. Koch (1940—), German for
“cook,” “coctor” in Latin, one of the collectors of the type material and most
valued friend of Botany and botanists in México

ained on at tne, A. Harriman, o nov. (Fig. 2). Type: MISAC O.
MicHoacan: Tizupan, Coalcoman, woods, 0.5 m high, flower red, alt. 50 m, 4 Sep
oa ae et 7 15917 (noLoryPeE: LL!; isorypes: K, NY, US and een

A b li landul fl fere uncinatis | | lal 1
: oOo f oO

Perennial the slender stems probably sprawling, to 0.5 m tall or
long, thinly to densely eglandular hairy, the hairs hooked and appearing
almost uncinate, antrorsely inclined. Leaves opposite, isophyllous, blades
25-40 mm x 15-25 mm, ovate, apically retuse or rounded or acute, ba-
sally cuneate to rounded, with evident internal crystals, indistinctly cili-
ate, with a few hairs on the veins abaxially, otherwise glabrous, on hairy

682 Sipa 18(3)

Treated in 1967 with
‘aury! pentachlorphenate!

<p Blela

-

ey ey 9) Ip Pe
as

id
,Ojr E je)
1,
See

a

LUNDELL
HERBARIUM

}

ep yt

:
j

versity of Texa
AUSTIN, TEXAS

—

jp OR Ait, Sty Sit wt | eit

: ICAL GARDEN
abs, otte@ appellee,
f Pprehased. tga
vemeaN FSP and~ cl
elele rig panicula 0
tett GAT Re Cie

EPO BET cnpi

foodus , ose hag t yea; Leoales

Shetpn w tet Shots. : Joal-

coman, Woh,, “ex,, ool), He et wl.
det. Morton

Fic. 2. Holotype of Commicarpus praetermissus N.A. Harriman.

Harriman, New World Commicarpus 683

petioles 2—4 mm long. Flowering branches from the upper axils and termi-
nal, bearing numercus pedunculate umbels, the peduncles and pedicels glabrous.
Perianth above the ovary red, funnelform, 9 mm high and 12 mm broad,
glabrous; stamens 2, the anthers exserted on glabrous filaments; style ex-
serted, stigma capitate; fruit (anthocarp) slendetly clavate, 7.5—9 mm long,
striate, glabrous, the prominent glands in 3—5 well-defined horizontal bands.

Flowering and fruiting.—September, so far as known.

Distribution and habitat.—Known solely from the type collection; from
the Pacific coast at 18°11' N, 103°2' W (Hinton & Rzedowski 1972

Etymology.—The epithet refers to the fact that the specimens have lain
overlooked and neglected in numerous herbaria for over half a century; C.V.
Morton in preparing the labels remarked that it was a new species; Meikle
(1978) likewise remarked that it was a new species, though the material he
had at hand from K was insufficient for a proper description.

KEY TO THE NEW WORLD SPECIES OF COMMICARPUS

Perianth above the eee ae white or greenish-yellow, at most pink-
veine the sinuses; stam
Ze Penland white or ere 34 mm long and wide, externally glabrous
or obscurely puberulent; widespread from Texas to se Arizona and Baja
California, s to Guatemala; West Indies; Bahama Archipelago; Colombia
and Venezuela to Per Commicarpus scandens - ) Standley
2. Perianth white, eae veined in the sinuses, 10-15 mm long and 7
mm wide, hirtellous externally; endemic to Baja California, ee

ommicarpus brandegeei Standley
. Perianth above the developing fruit light purple to mauve to cherry red;
a ,3,or4
3. Stems and leaves glabrous; perianth lilac, pink, or purplish red, puberu-
lent at tip in bud, otherwise glabrous; stamens 3; Bolivia, Peru, Ecuador
and Galapagos Islands Com

eee

micarpus tuberosus (Lamarck)
Standley, to include C. oe Heimerl

punters and leaves variously hai red, stipitate glandular or glabrou
4. Stems viscid eeauay with spreading haiis: peraath abundantly ss
shortly ex ternally, to 16mm hi gh and broad; stamens

4; thus far known only from Oaxaca, México

a ae coctoris
N. arriman
4. Stems with abundant short, non-glandular hairs, these hooked at ce
tip and antrorsely inclined; perianth glabrous externally, to 9 mm high
and 11 mm broad; stamens 2; thus far known only from Michoacan,
Co

México mmicarpus praetermissus N.A. Harriman

ACKNOWLEDGMENTS

Paul A. Fryxell, now “retired” to TEX, has been a continuous source of
encouragement, help, and editorial acumen.

684 Sipa 18(3)
REFERENCES

Acevebo-Ropricuez, P. 1996. Flora of St. John, U.S. Virgin Islands. Mem. New York Bot.
Gard. 78:1-581.

Boater, A. LINN. 1974. The genera of Nyctaginaceae in the southeastern United States. J.
Arnold Arbor. 55:1—37

Brako, L. and J. ZaRuccHI. 1993. Catalogue of the flowering plants and gymnosperms of
Peru; Catalogo o las angiospermas y gimnospermas del Per. Monogr. Syst. Bot. Mis-
souri Bot. Gard. 45:1—1286

BREEDLOVE, D. en re Floristicas de México. IV. Flora de Chiapas. Instituto de Biologia,
UNAM, México.

FosperG, ER. 1978. Studies in the genus Boerhavia L. (Nyctaginaceae), 1-5. Smithsonian
Contr. Bot. 39:1—2¢

GONZALEZ ELIZONDO, M., $8. GONZALEZ ELIZONDO, and Y. HERRERA ARRIETA. 1991. Listados
Floristicos de México. IX. Flora de Durango. Instituto de Biologia, UNAM, México.

eet . and J. RzEpowsk1. 1972. as B. Hinton, collector of plants in southwest-

xico. J. Arnold Arbor. 53:141-181.

Poe R. 1988. Flora of the am ree 4:1-673.

Liocier, H. 1985. Descriptive flora of Puerto Rico and adjacent islands. 1:1—352.
EIKLE, R.D. 1978. A key to Commicarpus. Notes Roy. Bot. Gard. Edinburgh 36:235-249.

STANDLEY, P.C. 1909. The Allioniaceae of the United States with notes on Mexican species.
Contr. U.S. Natl. Herb. 12:303-389.

__. 1911, The Allioniaceae of Mexico and Central America. Contr. U.S. Natl.

‘He i 13:377-280.

MUHALENBERGIA JAIME-HINTONIT
(POACEAE: CHLORIDOIDEAE),
A NEW SPECIES FROM NUEVO LEON, MEXICO

PAUL M. PETERSON

Department of Botany
National Museum of Natural History
Smithsonian Institution

Washington, DC 20560-0166
JESUS VALDES-REYNA

Departamento de Boténica
Universidad Autonoma Agraria “Antonio Narro”
Buenavista, Saltillo
Coahuila 25315, MEXICO

ABSTRACT

Mubhlenbergia aie -hintonii P.M. Peterson & Valdés-Reyna, sp. nov., is described and il-
lustrated. The new speci es occurs in gypsum soils in southern Nuevo Leén, Mexico be-
ween the municipalities of Aramberri and General Zaragoza. Based on anatomical and
morphological features Muhlenbergia jaime-hintonii is placed in subgenus Podosemum, sec-
tion ies The new species seems most closely allied with Pee pubighuma buat
differs by its shorter culms (44-82 cm oe shorter truncate ligules (0.4-1.0 mm mee
with ciliolate apices, shorter spikelets (1.5—2.1 mm lon ng), shorter glumes (0.7—1.2
long), shorter lemmas (1.5—2.0 mm long), Re paleas (1.5—2.0 mm long), and ae

anthers (0.8—1.0 mm long).

RESUMEN

Se describe y se ilustra Mublenbergia jaime-hintonii P.M. Peterson & Valdés-Reyna, sp.
nov. Esta nueva especie se presenta en pa rte de las tierras gips6filas en el sur del estado de

Nuevo Leon, México, en | y General Zaragoza. Basados en caracteristicas

anatémicas y morfolégicas Mublenber oe jaime- saasiiie se ubice en el Supzeneho) icisiliniia
secci6n Epicampes. Esta nueva especie est

pero difeie a ella por sus ne mas cortos ee 82 cm de a ligula mas cortas
(0.4—1.0 mm de largo) trunc 4 lados, espiguillas mas cortas (1.5—2.1 mm
de largo), glumas mas cortas (0.7—1.2 mm de largo), lemas mas cortas (1.5— - mm de
largo), paleas mas cortas (1.5—2.0 mm de largo), y anteras mas cortas (0.8—1.0 mm de largo).

While working on a catalogue of the grasses of Northeastern México (Valdés-
Reyna, Davila, & Carranza Pérez, in prep.) and while preparing a revision
of the perennial Muhlenbergia for México (Peterson, in prep.), JVR suggested
to PMP that a collection by Mr. Jaime Hinton from southern Nuevo Leén
might represent an undescribed species. At first glance the lemmas of this

Sipa 18(3): 685-691. 1999

686 Stipa 18(3)

specimen appeared to be 1-veined and the ligule appeared to be a line of
hairs, both characteristics of Sporobolus. Two specimens (Hinton 22098 & Hinton
3766) were previously sent to J.R. Reeder (ARIZ) who determined them
to be Sporobolus eryptandrus (Yorr.) A. Gray. However, upon closer inspec-
tion, the lemmas are usually 3-veined with faint lateral veins, these occa-
sionally absent; and the ligule is very short with a membranous base and a
truncate-ciliolate apex. The new species is clearly a member of the subfam-
ily Chloridoideae, tribe Eragrostideae (Peterson et al. 1995, 1997). There-
fore, we describe these specimens as a new species of Muhlenbergia, and name
the species after the original collector, Jaime Hinton.

eae eae jaime- sec P.M. Peterson & Valdés-Reyna, sp. nov. (Figs.
2). Type: MEXICO. Nuevo Leon: Municipio General Zaragoza: La Joya, Cuesta
Bienen: sexi 15 km S of i on road towards Zaragoza (23° 59! 37.1"
N-99 ° 47! 38.7" W), 1345 m, 31 Oct 1998, J. Valdés-Reyna & M. A. Carranza eee

2560 ona ANSM!; IsoTYPE: oo

=

A Muhlenbergia pubigluma en sueculs oe en aets apigomarieus aes folits,
igulis 0.4—

0.5—6.0 cm longis, spiculis 1.5—2. - mm ee Sa: 7—1.2 mm longis glabris, lemmatibus
|

culmis 44—82 cm altis, |

1.5—2.0 mm longis leviter 3(1)-venis, paleis 1.5—2.0 mm longis, antheris 0.8—1.0 mm longis,
recedit.

Loosely caespitose perennial with short, densely leafy rhizomes and ex-
travaginal shoot initiation. Culms 44—82 cm tall, erect, compressed keeled
near the base, densely white pubescence below the basal nodes, these hid-
den beneath the leafy sheaths, the hairs 0.8—1.3 mm long, upper nodes glabrous
or puberulent; internodes puberulent below and mostly glabrous above. Sheaths
6-28 cm long, longer than the lower internode, puberulent to glabrous,
stiff and brownish below, often curled; margins mostly smooth with a few
short hairs near the summit. — 0.4—1 mm long, membranous below;
apex truncate, ciliolate. Blades 5—22 cm long, 1.8—5 mm wide, flat just
above ligule to tightly conduplicate above, apically acuminate, Soriewhat
stiff, pubescent above and glabrous below. Panicles 13—34 cm long, 0.7—7
cm wide, narrow to somewhat open, the ascending densely flowered branches
tightly appressed or loosely spreading up to 40° from the culm axis; pedicels
0.5—2 mm long, ascending, scaberulous; inflorescence branches 0.5—6.5 cm
long. Spikelets 1.5—2.1 mm long, appressed to branches, 1-flowered, red-
dish-gray. Glumes 0.7—1.2 mm long, oblong, shorter than the lemma, usually
equal in length, faintly 1-veined, reddish, glabrous, scaberulous along the
midvein; apex acute to obtuse, occasionally minutely erose. Lemmas 1.5—
2.0 mm long, oblong to elliptic, unawned, faintly 3(1)-veined, greenish
mottled with reddish areas; midvein, margins, and proximal 1/2 to 3/4 loosely
to densely appressed pubescent to villous, often these hairs more numerous
along the margins and midvein below, the hairs up to 0.5 mm long; apex

PETERSON AND VaALDEs, Muhlenbergia jaime-hintonii 687

Fic. 1. Muhlenbergia jaime-hintonii (Valdés-Reyna & Carranza Pérez 2560). A. Habit. B. Sheath,
ligule, and portion of a blade. C. Inflorescence. D. Secondary branch of inflorescence. E.
Spikelet. F. Glumes. G. Lemma, dorsal view. H. Lemma, opened dorsal view. I. Lemma,
lateral view. J. Palea. K. Palea with stamens, pistil, and lodicules. L. Stamens, pistil, and
lodicules. Scale for C is shown in A; scale for F—L is shown in E.

688 Stipa 18)

acute, rarely minutely mucronate. Paleas 1.5—2.0 mm long, oblong, 2-veined,
equal in length to the lemma, the proximal 2/3 to 3/4 densely appressed
pubescent to villous between the veins and along the margins; apex acute
to obtuse. Stamens 3; anthers 0.8—1.0 mm long, reddish at maturity, greenish
when immature. Ovary 0.2 mm long; styles 2, separate, glabrous; stigmas
2, feathery, whitish. Caryopsis not seen.

Phenology.

g in October through November

site Nia Voi ri jaime-hintonii is known only from southern
Nuevo Leon between | 300-1850 m in the Municipio’s Aramberri and General
Zaragoza, and can be found growing in whitish, alkaline soils derived from
gypsum with Lewcophy/lum hintoniorum G.L. Nesom, Scutellaria lutilabia T.M.
Lane & G.L. Nesom, Galinm dempsterae B.L. Turner, Lobelia gypsophila T.J.
Ayers, Geniostemon eypsophilum B.L. Turner, Ca/fisia hintoniorum B. L. Turner,
Agave striata Zucc., and Hechtia glomerata Zucc.

Additional specimens examined: MEXICO. Mpio. Aramberri: Between La Escondida
and Aramberri, 1425 m, 23 Oct 1993, G.B. Hinton et al, 23707 (ANSM, US); San Fran-
cisco, 1835 m, 3 Sep 1993, G.B. Hinton 22698 (ANSM, TEX); Mpio. General Zaragoza:
Aramberri-El Salitre, 1325 m, 26 Oct 1993, G.B. Hinton 23766 (ANSM, TEX); La Joya,
Cuesta Blanca, approximately 15 km S$ of Ar. amibert rion road towards Gral. I Zaragoza 1345m,
29 Jul 1998, M.A. Carranza Pérez & J. Valdés-Reyna C-2981 (ANSM

LEAF ANATOMY

Cross-sectional leaf blade anatomy was determined from hand sections
of fresh field collected material (Va/dés-Reyna & Carranza Pérez 2560) on tem-
porary slides. The sections were stained in 0.05% toluidine blue. A dia-
grammatic cross-section was made with the aid of a camera lucida (Fig. 2).

The blades (Fig. 2) are typically kranz-C4, NAD-me (nicotinamide ad-
enine dinucleotide co-factor malic enzyme)-like in Hattersley and Watson’s
(1992) sense with tightly radiate arranged chlorenchymaand XyMS-+ (presence
of cells between the metaxylem vessel elements and laterally adjacent
chlorenchymatous tissue, see also Hattersley and Watson 1976). The lamina
are flat or conduplicate with primary and secondary vascular bundles about
the same size. The primary vascular bundles are well differentiated into xylem
with metaxylem, phloem, and a double bundle sheath (mestome and pa-
renchyma bundle sheath). The phloem tissue is irregularly sclerosed. The
ribs are flattened with angled sides (rectangular) and the furrows are 1/5 to
1/2 as deep as the width of the blade adaxially and usually less than 1/5 as
deep abaxially. The medium vascular bundle structure consists of a simple

eel with only a single primary vascular bundle. Per blade there are 9-15
primary vascular bundles and 16-24 secondary vascular bundles. There is
one or two secondary or tertiary vascular bundles placed between each pri-
mary vascular bundle. All vascular bundles are situated in the median layer

PETERSON AND VALDES, Muhlenbergia jaime-hintonii 689

Ce 2
Kk (-( 4)

a Sts
||

pbs

ms

As 4

RRS
Rare

A
bey

2
i-J
4)

Fic. 2. Leaf blade cross section of a primary vascular bundle (center) and two secondary/
tertiary vascular bundles of Muhlenbergia jaime-hintonii (Valdés-Reyna & Carranza Pérez 2560).

Scale bar = 25 (m; chl = chlorenchyma tissue; f = sclerenchyma fibers; ms = mestome sheath;

mx = metaxylem vessel; pbs = parenchyma bundle sheath.

of the blade and are rectangular in outline. The xylem of the primary vas-
cular bundles contains two wide metaxylem vessels that are about the same
size as the parenchyma bundle sheath cells. The mestome, or inner sheath,
is always present in the vascular bundles surrounding the xylem and ph-
loem. The mestome is composed of small cells with thick walls and in pri-
mary, secondary and tertiary bundles it is interrupted abaxially and adaxially
by sclerenchyma fibers forming a girder. The parenchyma bundle sheath,
or outer layer, contains large, thin walled cells that are interrupted by the
abaxial and adaxial sclerenchyma girder in the primary, secondary, and ter-
tiary bundles. Chlorenchyma cells radiate just outside the parenchyma bundle
sheath cells and are interrupted by a column of colorless cells forming a
tightly radiate arrangement (NAD-me-like). Five to eight rows of scleren-
chyma fibers form the abaxial and adaxial girders which are wide near the
epidermis and narrow toward the vascular bundle. A few fibers form a rect-
angular to round cap along the margin of blade. Thick-walled colorless cells
form a continuous column separating each vascular unit, these are not dif-
ferentiated into bulliform cells.

DISCUSSION

The new species seems most closely allied to species of subgenus Podosemum,
section Epicampes since it has culms with compressed keeled bases, condu-
plicate blades, primary and secondary vascular bundles both about the same
size with sclerosed phloem, and shallow adaxial furrows (Peterson, in press;
Peterson & Herrera-Arrieta, in review). Muhlenbergia jaime-hintonii differs
from M. emersleyi and M. pubigluma (see Table 1) by quite a few characteris

690

Sipa 184)

TaBLe |. Salient features comparing Muhlenbergia jaime-hintonii with M., emersleyi and M. pubigluma.

Characters

M. jaime-hintonii

M. emersleyi

M. pubiglhima

Shoot initiation
Rhizomes
Culm herght

Ligule length
(mid-culm)

Ligile apex
Inflorescence

I® branch length
Spikelet length
Glume length

Glume surface

extravaginal
short-leafy
44—82 cm

0.4—1 mm

truncate, ciliolate

0.5—6.5 cm

1.5-—2.1 mm
0.7—1.2 mm

glabrous

intravaginal

absent
100—150(—200) cm
|

J)—25 mm

acuminate, lacerate

(1—)9-17 cm

i)

2.2—3.2 mm

~)

2.2—3.2 mm

glabrous

intravaginal
short-non leafy
75-125 cm

5-13 mm
acuminate, lacerate

4—S cm

2.5-—3.5 mm
2.5-3.5 mm

pubescent

Lemma length 1.5-2.0 mm 2-3 mm 2.9—3.1 mm
Lemma veins faintly 3(1)-veined prominently 3-veined faintly 3-veined
Lemma awn absent present or absent present

Palea length 1.5-2.0 mm 1.8-2.9 mm 2.3-3.3 mm
Anther length 0.8—1.0 mm 1.2-1.6 mm 1.4-1.8 mm

tics, most notably: extravaginal shoot initiation, the presence of short-leaty
rhizomes, short ligules (0.5—1 mm long) with a truncate and ciliolate apex,
spikelets ranging from 1.5—2.1 mm long, short glumes (0.7—1.2 mm long)
that are glabrous, short lemmas (1.5—2.0 mm long) that are faintly 3(1)-
veined, short paleas (1.5—2.0 mm long), and short anthers (0.8—1.0 mm
long). Mahlenbergia distans Swallen is also morphologically similar to M.

jaime-hintonii but differs by having longer ligules (usually S—10 mm long)
that are firm below, longer spikelets (2.7—3.6 mm long), and short-awned
(the awn 1.5—5 mm long) lemmas (Soderstrom 1967). The closest sister to
M. jaime-hintonii appears to be M. pubigluma since they both share several
characteristics, such as: densely white pubescence below the basal nodes,
short-leafy rhizomes, reddish-gray spikelets, faintly 3-veined lemmas that
are pubescent to villous on the lower 1/2 to 3/4, and simple keels (com-
pound keels are the predominant character state in sect. Epicampes). In ad-
dition, Muhlenbergia pubigluma and M. jaimie-hintonii appear to be sympat-
ric since the former species is known to occur just 10 km west of Aramberri
between La Escondida and La Soledad [A.A. Beetle M-406 (UC, US!)}, and
again west of Doctor Arroyo, approximately 60 km southwest of Zaragoza

{Shreve & Tinkham 9651 (GH)Y}

PETERSON AND VALDES, Muhlenbergia jaime-hintonii 691
ACKNOWLEDGMENTS

Appreciation is extended to Alice R. Tangerini for providing the illus-
tration and Dan H. Nicolson for correcting the Latin diagnosis and discus-
sions pertinent to choosing the specific epithet. Stephan L. Hatch and Robert

. Webster are thanked for reviewing the manuscript on short notice. Miguel
A. Carranza Pérez is thanked for his help in obtaining the field collections.

REFERENCES
Harrerstey, P.W. and L. Watson. 1976. C4 grasses: An anatomical criterion for distin-
guishing between NADP- ne ‘ue species and PCK or NAD-malic enzyme spe-
cies. Austra. J. Bot. 24:297—
oa 1992. Diversification of photosynthesis. In: C.P.

Chapman, ed. ae evolution and domestication. Cambridge University Press, Cam-
bridge. Pp. 38-116.
PrTERSON, P.M. 199_. Systematics of the Muhlenbergiinae (Chloridoideae: Eragrostideae).
Proceedings of the Monocots II and Grasses III Conference (in press).
______and Y. Herrera-Arrieta. An anatomical survey of Muhlenbergia (Poaceae:
erence Syst. Bot. (in review).
_______, R.W. Werstrer, and J. Vatpfs-RryNa. 1995. Subtribal classification of
the i World Eragrostideae (Poaceae: Chloridoideae). Sida 16:529-544.
Eaereeesees ater ares —s 1997. Genera a - World
Braptostidese Coe Chloridoideae. Cation Contr. Bot. 87:

—

SODERSTROM, T.R. 1967. Taxonomic study of subgenus istiaibecis and section cea of
Mubhlenbergia (Giese. Contr. U.S. Natl. Herb. 34:75-189.

692 Sipa 18(3)
BOOK NOTICE

RICHARD RaBELER. 1998. Gleason’s Plants of Michigan: A Field Guide.
(ISBN 0-966325 1-0-9, pbk) Oakleaf Press, 920 Vesper Road, Ann Arbor,

MI 48103. 734-668-8579. $ 21.95. 398 Pp.
This book is a bargain. Don’t miss it !! After a nice introduction, a preface with a short
biographical sketch of Henry Allan Gleason, and a crystal-clear explanation of how to use
the book, Rabeler presents a nice bibliography, a very simple phytogr raphy section, a list of
illustrations, and then keys to the major groups of “plants” in the flora. The groups of
plants in the keys include woody plants, unusual plants, monocots and dicots. While these
are not the groupings one would normally encounter, they are, nonetheless, extremely use-

ful, especially for the amateur or hobbyist.

The book contains 49 pages of bracketed keys ightforward, simple, and nomenclaturally

harmonious with the magnificent technical eels Flora volumes by the other temper-
ate floristician at MICH, Dr. Ed Voss. Starting on page 93, there are descriptions for each
family, and bracketed keys to the species within each family (ordered so the species of each
lustrations, judiciously cho-

genus are adjacent to each other). There are 45 simple line i
sen, an 11 page glossary, and finally, indices to subjects and plant names. There are also
supplementary boxes scattered throughout the book, with extra information that the en-
thusiasc will appreciate, such as that found on page 299 for Galium (Rubiaceae), the bed-
straw, and for the infamous genus So/idago on page 331. These boxes explain points such as
the difficulty of the group, recent changes in taxonomic concepts and other information
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The cover is extremely attractive and sturdy, as is the binding. The paper is of high quality
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cesstul instructor for some time and has a knack for keeping someone interested even when
keying out a difficult group. He is to be congratulated for revising Gleason’s work and
producing a field manual ee with the Voss magna opt. The practicality, simplicity
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-John J. Pipoly 1

area near the Great Lakes Region!

Stipa 18(3): 692. 1999

A NEW SPECIES OF EMORYA (BUDDLEJACEAE)

MARK H. MAYFIELD!

Plant Resources Center
University of Texas
Austin, TX 78713, U.S.A.

ABSTRACT

A new species of Buddlejaceae, Emons ‘nconensts, is described here as the second species
of the genus. The species is known from a single collection from the Serranfas del Burro in
northern Coahuila, Mexico. ica a associated with a more mesic flora than Emorya suav eolens,

morphological adaptations and associated species suggest that E. rinconensis may grec
edaphically xeric, steep rock outcrops. The racemose inflorescences of the new species are
unique eee New World Buddigaceae and appear to represent a reduction from the open,
few-flowered axillary cymes found in Emorya suaveolens. In the context of North American
as in open inflorescences, long styles, linear calyx lobes,

Buddlejaceae, the long tubular corol
and prolate pollen support the present generic placement of the new species with Emorya,
t furure confirmation of the phyletic position of both species in a larger geographic con-

text is needed.

RESUMEN

En este trabaj lescribe una nueva especie de Buddlejaceae, Emory

la segunda especie del género. La especie se conoce a partir de una sola ele ta ade ii Seats
del Ba urro en el norte de Coahuila, México. Aunque asociada con una flora mds mésica que
Emorya suaveolens, las z Sa aaa morfol6gicas y jue E. rinconensis
puede crecer en afloramientos rocosos, escatpados y xéricos. Las infleteveeacins racemosas
de la nueva ae son tinicas entre las Buddlejaceae del Nuevo Mundo y parecen representar
una reduccién de las cimas axilares abiertas de pocas flores encontradas en E. swaveolens. En
el contexto de las Buddlejaceae Norteamericanas, las largas corolas tubulares en inflorescencias
ele estilos largos, Isbul os del caliz lineares, y polen prolado, apoyan la presente posicién
genérica de la nueva especie con Emorya, pero se necesita una futura confirmacién de la
situacion filética de ambas especies en un contexto geografico mas amplio.

A set of plant specimens from northern Coahuila, Mexico? distributed
to TEX for identification included an undescribed species which closely resembles
Emorya suaveolens Torr. of the Buddlejaceae (f. 1). The stellate-tomentose vestiture,
superior ovary, and four-merous, tubular corolla of the new species are characteristic
in Buddlejaceae, a family with only two genera in the New World: Emorya
and Buddleja (excluding Polypremum and Peltanthera which are better treated

'Current address: Herbarium, Department of Biological Sciences, Louisiana State Univer-
sity, Baton Rouge, — 70803, U.S.

Plants collected b SU Aguilar Ruiz: in association with a study of populations of Black
Bears in northern Coahuila, Mexico, at the Rancho E! Rincén. The first set of specimens of

the large collection from this study is at SRSC.

Stipa 18(3): 693-699. 1999

694 Sipa 18(3)

outside of Buddlejaceae, [Eliane Norman, pers. comm.}). Its racemose
inflorescences with solitary flowers in the upper axils are unique in the family
in the New World and appear to represent a reduction from the open cy-
mose axillary flowers of E. swaveolens. The species has not been collected again
since its initial collection in 1992, and the present collection is lacking fruits.
Thus, although fruits are expected to corroborate the present placement in

Emorya, the species is formally described here to advance its rediscovery and

thereby facilitate future research on the genus.

Emorya rinconensis Mayfield, sp. nov. (Fig. 1). Type: MEXICO: Municipio
Villa Acuna, cae | Rincon, SW s del Burro, in Cafion El Becerro,
28° 40'N 102° 15' W, 8 Aug oe Sholay Ruiz 164 with D.L. Doan-Crider (HO-
LOTYPE: ME an ISOTYPES: SRSC, TEX).

Valde diftert a E eles heaiosinied Torr. caulibus Senne aes eOupus e foliorum summorum
: ane

axillis solitaribus. cc tis multum densius vestitis

inter pilosis, staminibus se. ae VIX EXSEFTIS, volline eens

Spreading subshrubs with a conspicuous dense vestiture of 4-branched,
stellate trichomes on the young stems, leaves, and flowers. Stems ascending
distally to ca. 25 cm high; older woody branches to ca. 3 mm thick, witha
light gray exfoliating bark; young growth of stems ca. 1.2 mm thick and
densely stellate tomentulose. Leaves to ca. 2 cm long, opposite, estipulate,
the blades broadly oblanceolate to subspatulate or subrhombic, essentially
sessile or with a brief petiole; apices obtuse to rounded, with | to 4 broad
teeth along the margins; bases attenuate and entire; leaf surfaces densely
stellate tomentulose below, subglabrous above. F/owers solitary in the axils
of the upper leaves (i. e., paired at each node); pedicels ca. 4 mm long, the
pedicels each with pair of opposite, linear-elliptic, ascending bracteoles ca.
7 mm long closely subtending the calyx. Ca/yx 7.0—-7.5 mm long, narrowly
campanuliform with four erect to slightly spreading linear lobes 2.2—3.7
mm long, the tube 3.8—4.5 mm long, tapered to the base, with weak nerves
along the midribs. Coro//a 27-30 mm long, tubular, four-lobed, cinnabar-
ine, the color externally obscured by the vestiture of stellate trichomes; tube
ca. 1.2 mm wide at the base, scarcely expanded to the base of a short ampliate
throat which is constricted at the base of the lobes; lobes valvate in bud,
rounded-obtuse, spreading upwards at anthesis to ca. 4 mm across the top,
with scattered orange capitate resiniferous glands within; the interior of
the tube brick red, with a scattered pilosity of delicate, simple, minutely
papillose orange trichomes to 0.3—1.2{[—1.5] mm long. Stamens included within
the throat or slightly surpassing in late anthesis; filaments ca. 2.0 mm long,
inserted ca. four-fifths of the way up the tube (22—23 mm above the base);
anthers 2 mm long, introrse, mucronate at the apex, the anther sacs sepa-
rate below the attachment. Ovary narrowly oblong-elliptic, ca 4 mm long,
glabrous on the lower third, stellate-pubescent above, borne on a short

MayeleELb, A new species of Emorya 695

eae aoe
oe

t
fi
i
Tera
es
\
Lis
bo
t
i
bee
i
i
his
f
n

Fic. 1: A-D, Emorya rinconensis, from the type collection, Aguilar Ruiz 164. A. Branch showing
probable habit B. Flowering bes nch; C. Flower with pedicel and subtending bracteoles; D.
Corolla showing filament insertion; E. eee branch of Emorya suaveolens for compari-
son (from Nesom 7379).

stipe-like base ca 1.5 mm long; nectar disc present but inconspicuous; style
30-35 mm long, scarcely to evidently exserted from the corolla tube; stigma
truncate, scarcely bilobate, the surface papillose. Fraits unknown. Pollen prolate,
tricolporate, 31.5 pm x 18.5 pm
Distribution.—The species is as yet known only from the type collection.
Emorya rinconensis is associated with a flora similar to that of the Edwards
Plateau area of Texas. Its xeromorphic features are similar to other petrophilous

696 SipaA 18(3)

species of Buddlejaceae (e. g., Buddleja racemosa, of central Texas), which
inhabit dry limestone outcrops, often pendulous from steep slopes or ledges.
Other potentially rock-outcrop-dwelling species collected by Ruiz at the
type locality (“Cafion ae ae ude A geratina potosina, Desmodium lindhermert,
Eucnide bartonioides, Geranium caespitosum, Hedeoma costatum, Oenothera macrosceles,
Orbexilum ene Pion. Vlas Polygala scoparioides, Salvia regla,
Salvia roemeriana, Solidago nemoralis, Solidago petiolaris, and Thelesperma simplicifolium.
Future attempts to relocate Emorya rinconensis should be concentrated in areas

with limestone faces or ledges having these species.

Emorya suaveolens Torr. is distributed in the northern Mexican states of
Nuevo Le6n and Coahuila and in the United States along the Rio Grande
in Closely adjacent Texas (Fig. 2; Norman & Moore 1968; Norman 1964;
Warnock 1964). It is a shrub or small tree to 2 m tall with flowers in open
thyrses, with tubular, yellow corollas, exserted stamens with the filaments
inserted on the lower half of the tube, deltoid leaves, and tetracolporate pollen
(Norman & Moore 1968; Fig. 3). In contrast, E. rinconensis is a small spreading
shrub with reddish-orange flowers in racemes, filaments inserted supramedially
on the tube, included stamens, obovate to subspatulate leaves, and tricolporate
pollen (Fig. 3b). These species share a combination of features that separate
them from Buddleja in North America including: long tubular corollas, prolate
pollen (Fig. 3), anthers on elongated filaments (vs. subsessile), linear bracteoles
on the peduncles, linear we lobes, hypogynous nectaries, and leaves with
repand margins and non-clasping bases.

The new species is somewhat intermediate between Emorya suaveolens and
some species of Buddleja. Indeed, authorities on Buddlejaceae and related
groups have suggested that E. svaveolens could be included within the ge-
nus Buddleja (e. g., Leenhouts 1963; Norman 1967; Rogers 1986). Despite
this, Emorya has never been formally transferred nor has any species or group
of species within Buddleja ever been cited as a potential close relative. At
least some lines of evidence suggest that Emorya may not be so closely re-
lated to Buddleja. Although viable hybrids between distantly separated species
of Buddlejaceae have been produced (e. ¢., Nicodemia madagascariensis [Madagascar]
x Buddleja globosa {Chile} by Van de Weyer [1920}), Norman & Moore (1968)
reported an inability to produce viable hybrids between E. swaveolens and
some species of Buddleja (notably, seeds and seedlings were produced in crosses
with Buddleja alternifolia {Asian} and B. tubiflora {South American], but the
seedlings died prior to elongation of the hypocotyl). Norman & Moore also
pointed out morphological, anatomical, and palynological features that suggest
that Emorya may be phyletically distinct from Buddleja in Mexico. Norman
(1967) has also indicated that most of the New World species of Buddleja
are functionally dioecious, whereas E. swaveolens has perfect flowers. Punt
and Leenhouts (1967) assigned a distinct pollen type to Emorya stating that

697

MayriELp, A new species of Emorya

30°05' N
104°10'W 100 km =

Coahuila

Nuevo Leon

oO ’
| 2425 N
oO '
9945 W
Fic. 2: Documented distribution of the genus Emorya based on specimens at TEX-LL. Emorya

suaveolens (@); E. rinconensis (a).

the pollen morphology “strongly supports” continued generic recognition
for the species. In 1980, Punt reaffirmed this position and emphasized the
large size of the grains (ca. 40 pm vs. 25 pm for Buddleja), the tetracolporate
exine, and unique features of the columella (rather than the prolate shape).

698 SIDA 18(3)

Fic. 3: A-B. Pollen (2070x) A. Pollen of Emorya suaveolens. B. Pollen of E. rinconensis.

Without mature fruits and knowledge of the reproductive biology of the
new species, its generic placement with Emorya will remain somewhat equivocal.
Thus, the description rendered here for the new species places it with Emorya,
the Buddlejaceous genus in North America to which it bears the greatest
similarity.
ACKNOWLEDGMENTS

I wish to thank Eliane M. Norman for her informative discussions and

patient support, the herbarium of Sul Ross State University (SRSC) for loaning

their material of Emorya rinconensis, Sidney F. and Doris Blake Fund for funding
for the illustration, John Mendenhall for his aid with SEM photography,

—

and the Cell Research Institute of the University of Texas for use of its fa-
cilities.

REFERENCES

Norman, E.M. 1964. Additional data concerning the distribution of Emor lens. SouthW.
Nat. 9:2006.
. 1967. The genus Buddleja in North America. Gentes Herb. 10:47-114.
and R.J. Moore. 1968. Notes on Emorya ( Loganiaceae). SouchW. Nat.
13:137-142.
Punt, W. 1980. Pollen morphology. In: Leeuwenberg, A.J.M., ed. Die natiirlichen
PHanzenfamilien, Loganiaceae, 28b(1). 28b(1):162—191. Duncker & Humbolt.
—_ and P.W. PeeHOUr: 1967. Pollen Morphology and taxonomy in the
‘6 oganiaceae. Grana Palynol. 7:469—5 16

Rocers, G.K. 1986. The genera af ee in the southeastern United States. J. Arnold
Arbor. 67:143—185

SMITH, L.B., E.F. Guimaraes, J.E. Pereira, and E.M. Norman. 1976. Loganiaceas. In: Reitz,
P.R. (ed.). Flora ilustrada Catarinense I. Itayai, Santa Catarina, Brasil. Herbario Barbosa
Rodrigues. 1-77

MayrfiELb, A new species of Emorya 699

Torrey, J. 1859. Botany of the boundary. V. 2, p. 1. In: Emory, W.H. Rep. US and Mex.
Bound. Surv. Washington. A.O.P. Nicholson.

WeyER, WAN bE. 1920. Hybrid Buddlejas. Gard. Chron. ser. 3, 68:181.

WARNOCK, B.H. 1964. Verification of occurrence of Emorya suaveolens in western Texas. South W.
Nat. 9:206.

YOuNGMAN, A.L. 1964. Emorya (Loganiaceae) a member of the Texas flora? SouthW. Nat.
8:210-212.

700 Sipa 18()
BOOK RECEIVED
Kucera, CLair L. 1998. The Grasses of Missouri. Revised Edition. ISBN
Q-8262-1164-X, pbk). University of Missouri Press, 2910 LeMone
Boulevard, Columbia, MO 65201. 573-882-0180, fax 573-884-4498.
$34.95. 305 Pp. 269 Line drawings.

The Grasses of Missouri was first published in 1961. Significant changes have been made
in grass Classification since the first edition. Clair L. Kucera provides a complete and com-

prehensive classification of Missouri's vast grass flora. From the tallgrass prairies to
ested Ozarks to eae lowlands, the state offers an nareay of grasses that can be nae
fied into six subfamilie the Poaceae, eighteen tribes, and eighty-seven genera.

From the Cover: “Cover ring more than 275 species a flee entries, The Grasses of
Missouri is an essential research tool for identifying grasses, complete with working ke
descriptions, line drawings, distributions, a glossary, and a bibliography. The seolecsiondl
and lay person alike will benefit from this indispensable manual.”

Contents include:

Preface

Introduction

The Vegetation of Missc

Synoptic List of Missouri ‘Subfamilies, Tribes, and Genera

Key to Tribes

Keys to Genera

Description of Genera and Species

dssary
Bibliography
Index

Sipa 18(3): 700. 1999

DISCOVERY OF THE INDO-MALESIAN GENUS
HYMENANDRA (MY RSINACEAE)
IN THE NEOTROPICS,
AND ITS BOREOTROPICAL IMPLICATIONS

JOHN J. PIPOLY II

Botanical Research Institute of Texas
509 Pecan Street, Fort Mara TX 76102- 00 USA,
ppipoly @brit.org; clusia@latinmail.com

JON M. RICKETSON

Missouri Botanical Garden
P.O. Box 299, St. Louis MO 63166-0299, U.S.A.
jon.ricketson@mobot.org; jricketson @lehmann. mobot.org

ABSTRACT

taxonomic revision of a neotropical group of taxa formerly placed by Lundell in
Auriculardista, Chontalesia, and lcacorea, resulted in the discovery that they were actually
me

herwise Indo-Malesian genus, Hymenandra. The genus is defined by its
filaments fused at least basally to ae a staminal tube, the tube adnate to the base of the
corolla tube and anthers connate along their poe dehiscence sutures at least until
anthesis. It is also noted that all species of the genus exhibit Fagerlind’s Architectural Model,

a model that is otherwise extremely peccnicced, and one whose vegetative vs. reproductive
shoot dimorphism has led t to taxonomic Sipe aoe Nine species of Hymenandra are
recognized in the Neotropics, bringing the total number of species in the genus to 16.
Owing toa lack Fae Bea for Indo- . species, and see workers’ crea
to pneduivocaly aioe all of those apesiss toa subpcus: the neotropical group
subgenera. It is po. Paioe that the ae
of Hy nena dee. now considered an amphi taxon, is consistent with that predicted by
the Boreotropics Hypothesis, and its logical extension postulated by Wendt (1993) and
Lavin and Luckow (1993), especially when viewed in light of the biogeography for the rest
of the tribe Ardisieae. Each neotropical Hymenandra species is fully described, newly illus-
trated, and provided with citations for all taxonomic and nomenclatural synonyms. In addition

to the general section on morphology, biogeography and ecology provided for the genus,
each species is accompanied by a discussion of its distribution, ecology and conservation
status, along with citation of specimens examined. Five binomials are newly relegated to
synonymy, ace one taxon, a id pes is lectotypified. The following new combi-

’ onn. Sm.) Pipoly & Ricketson, H. ¢alycosa (Hemstl.)
Pipoly & Ricketson, H. sordida @rund ell) Dinky & Ricketson, H. sqvamata (Lundell) Pipoly
& Ricketson, H. w oe ae eee Pipoly & Ricketson, H. pittieri (Mez) Pipoly & Ricketson,
H. callejasii (Pipoly) Pipoly & Ricketson, H. acutissima (Cuatrec.) Pipoly & Ricketson, H.
crosbyi (Lundell) Bil & Ses A complete list of exsiccatae is provided at the end
for all collections examine

Stipa 18(3): 701-746. 1999

702 SIpA 18(3)
RESUMEN

Al llevar a cabo una revisi6n taxonémica de un grupo comprendiendo taxa anteriormente
clasificados por Lundell | los géneros Auriculardisia, Chontalesia, y Icacorea,

se revel que todos s eageiaaaeia al género indo- malestace, Hymenandra. E\ género se define

por sus filamen ados, formandose un tubo estaminal, por lo menos en su base, adnado

ala ne del tubo corolino y con anteras conadas a lo largo de las hendiduras longitudinales

de dehiscencia. Se note que todas las especies del género se caracterizan por exhibir el Modelo

Arqutectural de Fagerlind, un elo | tinico y cuyo dimort?: ramas vegetativas

y ae tivas se ha ocasionado sobredescripcion taxonomica. Se reconocen nueve espec
ales de

Hymenandra, samando el total para el género hasta 16. Debido ' ‘hecho 2.
que ry material indo-malesio es muy incompleto, no se han podido asignar a un subgénero

todas las especies. Por lo tanto, se transfierren las especies Bebra p ical aa sin circcumscribir
subgéneros. Se postula que la geografia del género esta de ac | patrén fitogeografico

proveniente del Hipotesis av se especialmente en vista de la distribucién y relaciones
el tribu Ardisieae. Ademas de la secci6n general sobre morfologia, biogeografia, y ecologia
proveida para el género, se ae la Aisabecibn geografica, ecologia y estado de conservacién,
se lista pliegos examinados para todos los taxa. Se relega cinco binomiales a la sinonfmia,
y se lectotipifica Hymenandra pittieri. Las nuevas combinaciones se incluyen a continuacioén:
The following new combinations are made: ett stenophylla (Donn. Sm.) Pipoly &
Ricketson, H. calycosa (Hemsl.) Pipoly & Ricketson, H. sordida (Lundell) Pipoly & Ricketson,
H. squamata (Lundell) Eipely & Ricker tson, H w iain (Lundell) Pipoly & Ricketson,
H, pittieri (Mez) Pipoly & Ricketson, H. ca/lejasiz (Pipoly) Pipoly & Ricketson, H. acutissima
(Cuatrec.) Pipoly & Ricketson, H. — (Lundell) Pipoly & Ricketson. Se provée una lista
completa de exsiccata al final del trabajo

INTRODUCTION

The pantropical genus A rdisia Sw. is by far the largest in the family Myrsinaceae,
containing perhaps as many as 500 species (Chen & Pipoly 1996). Its cir-
cumscription has been problematic owing toa lack of comprehensive treat-
ment since that of Mez (1902) in Engler’s Pflanzenreich, almost a century
ago. Several genera have been separated from Ardisia sensu lato since the Mez
monograph, including Chontalesia Lundell (1982), described as a monotypic
genus to accomodate the Central American Ardisia calycosa Hemsley. Ardisia
calycosa may be immediately recognized by its unique large, coriaceous, narrowly
lanceolate to lorate sepals that are longer than the corolla. While examin-
ing Ardisia calycosa to determine its relationships, we noted that its vegeta-
tive characters, such as the punctate and punctate-lineate, nitid leaves, the
branchlet apices with varying amounts of furfuraceous lepidote scales, and
most notably, the unusual leaf dimorphism between the floral and vegeta-
tive shoots (now known as an artifact of its morphogenetic program that
exhibits Fagerlind’s Architectural Model, sezsw Hallé et al. 1978), were similar
to a Central American suite of species that includes A. stenophylla Donn.
Sm., A. oblanceolata Standl., A. calvarioana Lundell, and A. wilburiana Lundell.
Careful study of this group a species revealed that their androecia have filaments
connate throughout their length to form a staminal tube (therefore mona-

—

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 703

delphous) basally adnate to the corolla, and anther thecae connate along
the suture line, at least in the basal portion, prior to anthesis, at which time
at least the distal (abaxial) portion of the theca opens to permit release of
pollen, or the anthers split completely apart from each other. This struc-
ture was also reported by Pipoly (199 2a) in the description of Ardisia callejasii
Pipoly, a species endemic to the Antioquian Chocé of northern Colombia
Careful reexamination of additional material of Ardisia callejasii has revea led
that the anthers are also basally connate up to full anthesis, when they split
apart completely. In addition, the combination of long-pedunculate panicles
terminating in very long-pedicelled flowers in corymbs to subumbels and
large fruits with thick, juicy exocarps common to all the aforementioned
taxa indicated that other neotropical species with the same features, including
Ardisia crosbyi Lundell, A. pittieri Mez, and A. acutissima (Cuatrec.) Lundell
might also belong to the alliance. Finally, with the sole exception of Ardisia
callejasi1, it was striking to note that for all species, the inflorescence was
borne terminally or subterminally (but behaving terminally) on the end of
a long sylleptic shoot, bearing leaves similar in shape to those of the veg-
etative shoots, except much smaller in size. The latter phenonmenon was
noted by Mez (1902) in Ardisia subgenus Pyrgus (Lour.) Mez, and by Stone
(1991) in the genus Hymenandra A. DC. ex Spach. All of the aforementioned
species except Ardisia calycosa have been referred by Lundell (1981) to his
genus Auriculardisia, defined principally by the asymmetric sepals auricu-
late in outline. While surveying the rest of the neotropical Ardisia species
attributed to Auriculardisia, we noted that Ardisia squamata (Lundell) J. E.
Morales and A. sordida (Lundell) J. F. Morales possess all the features of the
aforementioned group of taxa, but in a more diminutive form because they
are subshrubs.

Given that Ardisia has traditionally been separated from its close conge-

ers by filaments considered free from each other and from the corolla tube
(Giada 1966, 1971; Mez 1902; Ricketson & Pipoly 1997), the aforemen-
tioned group of species posed a serious problem in the circumscription of
what was admittedly already a parapyletic group (Stahl 1996). On the other
hand, members of Ardisia subgenus Graphardisia (Pipoly & Ricketson 1998)
have filaments basally fused for less than 1/5 their length, to form a short
staminal tube free from the corolla tube, that is inconspicous because of its
thinly membranaceous texture, and anthers that are totally free from each
other. If Ardisia is defined by stamens with free anthers and filaments free
or variously connate, but totally free from the corolla tube, the group of
species related to Ardisia calycosa discussed above would still not fit. The
inclusion of Ardisia calycosa and its relatives in Ardisia would significantly
amplify the morphological circumscription of the genus. To answer whether
further enlargement of Ardisia’s circumscription was warranted to main-

704 Sipa 1L8(4)

tain A. calycosa and its relatives relatives within Ardisia, a review of the re-
lated genera of the family was conducted. This led to the surprising con-
clusion that the relationships of this suite of species are clearly with the
genus Hymenandra, heretofore considered an Indo-Malesian genus of seven
species (Stone 1991).

THE GENUS HYMENANDRA

Hymenandra was described as a section of Ardisia by Alphonse de Candolle
(1834, 1841) to accomodate two species, A. hymenandra Wall. and A. glandulosa
Roxb., with filaments basally connate, but free apically, and anthers con-
nate. De Candolle (1841) treated the group as a genus, a rank to which it
had already been assigned by Spach (1840). Mez (1902) accepted the genus
in his worldwide monograph of the family, using the same characters. Furtado
(1958) transferred Ardisia iteophylla Ridley to the genus, on the basis of its
fused filaments and anthers, but in the same paper described A. calcicola
Furtado, distinguishing it from its sister species in subgenus Pyrgus by the
fused filaments and anthers. Nayar and Giri (1975) described one new spe-
cies, Hymenandra narayanaswamii Nayar & Giri, endemic to Myanmar (Burma).
Stone (1991) revised the genus, transferred A. calcicola to Hymenandra, and
described four new species, recognizing a total of seven species. Stone (1991)
erected two subgenera, based on the degree of compaction of the inflorescence,
the presence or absence of papillae on the adaxial calyx lobe surface, and
general habit of the plants. Hymenandra subgenus Lacrimophila B. C. Stone
was defined by caducous floral bracts, sepals hirtellous along the margins,
glabrous or hirtellous (not papillose) within, but he included only Hymenandra
narayanaswamii and H. diamphidia B. C. Stone in it, leaving the third spe-
cies of the group from the second half of his key, H. steophylla (Rid1.) Furtado,
in subgenus Hymenandra. We have examined material of both groups and
conclude that further material of the species will be necessary before ac-
cepting the subgeneric grouping.

Hymenandra, as defined by Stone (1991), is characterized by a staminal
tube formed by filaments at least basally united and basally adnate to the
corolla, concomitant with anthers connate, at least basally, by their longi-
tudinal dehiscence sutures, until anthesis, at which time the distal flap of
the theca may open or the anthers may split apart completely. All species
have filaments basally connate and apically free, with anthers connate in
anthesis along the entire length of the dehiscence sutures. In the Neotropics,
the filaments are connate throughout their length, at least at first, and the
anthers are connate in the basal half, separating early or just after the co-
rolla lobes open, whereas in the paleotropical species, the filaments are con-
nate in the basal half, free distally, and the anthers are connate along the

PipoLy AND RICKETSON, Hymenandra and its boreotropical implications rieb)

deshiscence sutures throughout their length. As far as is known, all mem-
bers of the genus exhibit Fagerlind’s Architectural Model, as defined by Hallé
et al. (1978). Given the uniqueness of the androecial structure, we here transfer
the aforementioned Ardisia species to Hymenandra.

With the addition of the neotropical species transferred in this paper,
the total number of species in Hymenandra reaches 16, distributed from Assam,
Bangladesh, Myanmar, Malaya, Borneo, and Nicaragua through the Colombian
Choco. Given the scarcity of material available for study, we defer reexami-
nation of subgeneric delimitation until a later time.

MORPHOLOGY

The following descriptions are based on the genus as a whole, with par-
ticular emphasis and detail among neotropical taxa.

Habit, Architecture, “Trunk” and Shoots

Most members of neotropical Hymenandra are shrubs or small trees to 8 m
tall with a maximum known diameter at breast height (dbh) of 8 cm, with
the exceptions of H. squamata and H. sordida, which are subshrubs less than
one meter tall. This is much like the case among the Indo-Malesian spe-
cies, where seven of the nine species are also subshrubs. Therefore, when we
refer to a “trunk” in the following architectural model discussion, it is the
principal axis of the plant, or “vegetative stem.” So far as is known, all spe-
cies of Hymenandra are terrestrial (i.e., not epiphytes).

Careful study of herbarium specimens and examination of photographs
lead us to postulate that all Hymenandra species exhibit Fagerlind’s Archi-
tectural Model (Hallé et al. 1978). Fagerlind’s Model describes those trees
whose architecture is determined by a monopodial, orthotropic, readily
distinguishable, rythmically growing trunk that produces tiers of modular
branches, each branch sympodial and plagiotropic by apposition, with Spi-
ral or decussate (never distichous) phyllotaxy. Branch modules are often
hapaxanthic (dying after flower) or, as is the case in Hymenandra, they may
exhibit sympodial growth by apposition, often branching several times without
flowering. Deducing evidence we have pieced together from specimens, field
observations, and collectors’ notes, we might explain the morphogenetic
program of Hymenandra individuals as follows: a) A “trunk” (“vegetative
shoot’) develops rythmically, producing successive pseudoverticels of “vegetative”
leaves until the apex loses dominance and latent lateral (“axillary”) buds
are released, producing successive, monopodial pseudoverticels of shoots,
each of which is sylleptic (and rarely attaining 5 mm in diameter), with a
long hypopodium and exhibiting rapid extension growth. b) The sylleptic
branch shoots produce first a prophyll, then pseudoverticels of “reproduc-
tive shoot” leaves (similar to the leaves of the “vegetative shoot” but nota-

706 SipA 18(3)

bly smaller), and then may either lose dominance and once again branch
sympodially by apposition growth (repeating the module) without flowering,
or produce a terminal or pseudoterminal (acting as a terminal) inflorescence.
Therefore, like the other species exhibiting Fagerlind’s Model, and unlike
other models, inflorescence production is usually not acropetal along a set
of modular plagiotropic branches (although it can be). c) When a shoot flowers,
the apex of the reproductive shoot will gradually lose dominance and may
produce the remaining flowers of the inflorescence, or support ongoing fruit
development, or the remaining flowers may abort. Following flower, a branch
sympodial by substitution growth may (or may not) be produced, that will
very slowly generate leaves in a high phyllotactic spiral (a pseudoverticel)
and eventually abort. It is interesting to note that the apical region of the
principal plane axis (“trunk”, or “vegetative shoot”) does not produce an
inflorescence, and as Hallé et al. (1978) noted, leaves associated with the
trunk (here termed “vegetative shoot leaves”) are considerably larger than
those of the branches (here termed “reproductive shoot leaves”). It 1s pre-
cisely this rare leaf dimorphism that caused us to further investigate the
dynamics associated with Fagerlind’s Model to see if they were consistent
with what we could surmise about the developmental biology (morphoge-
netic program) of Hymenandra species. Clearly, more intensive field studies
are needed to determine if any other activity occurs in the plant body’s de-
velopment among species of the genus.

Mez (1902) described the branching dynamics and flowering phenom-
ena associated with Fagerlind’s Model when he discussed the “inflorescences
on special branches, with the principal axis indeterminate, [the inflorescences]
terminal on lateral apices, with large subtending leaves subverticillate” for
Ardisia subgenus Pyrgus. Subgenus Pyrgus is Indo-Malesian in origin and
its species often occur sympatrically with individuals of Hymenandra. Like-
wise, Stone (1991) made the same comparison when he described Hymenandra
lilacina distinguishing between vegetative and flowering shoots in that species.
While both Mez and Stone noted that the leaves of the vegetative shoot
(crunk) are usually slightly different from those of the flowering shoot, it
was not correlated with a model for growth. During the period Mez worked,
the concept of architectural models did not exist, and the herbarium mate-
rial available to Mez was rather fragmentary or restricted to flowering shoots,
at times accompanied from a single leaf from the trunk or an axis in vegeta-
tive state. Stone (1991) noted similarities in the “position and form of the
inflorescence” between Hymenandra and Ardisia subgenus Pyrgus but did
not state this in terms of architectural models. While there are several other
architectural models present in Ardisia and within the tribe Ardisieae (Maasart’s,
Scarrone’s, Roux’s, based on unpubl. data), very few taxa have been exam-

Pipoty AND RICKETSON, Hymenandra and its boreotropical implications 707

ined, and architectural models are not necessarily correlated with subgeneric
or other taxonomic groups. While Hymenandra species may share the same
architectural model with Ardisia subgenus Pyrgus, the free anthers with apical
or subapical pores with minute filaments, and the numerous, pluriseriate
ovules of Pyrgws clearly preclude inclusion of the Hymenandra species within it.

Leaves

The leaves of Hymenandra are alternate, exstipulate and simple. As previ-
ously stated, leaves on the trunks (“stems” for small-statured taxa, “vegeta-
tive shoots” otherwise) are much larger than those of the reproductive ones,
and also have a smaller length-to-width ratio. While both the “trunk” (or
stem) and branches have spiral phyllotaxis, it is presumably by secondary
twisting of the branches that make them plagiotropic, rather than by dis-
tichous arrangement. The blades may be chartaceous to coriaceous, ellip-
tic, oblanceolate or oblong, apically acute to long-acuminate, basally cu-
neate to obtuse or rounded, decurrent on the petiole or not, conspicuously
or inconspicuously black punctate and punctate-lineate, usually glabrous,
but sometimes sparsely and minutely furfuraceous lepidote, the margins
are entire, or rarely crenate (Hymenandra calycosa). The petioles are margin-
ate and may bear the same tomentum as the branchlet apices, but they are
glabrescent,

Inflorescence, Flowers and Fruit

The inflorescence may be terminal or pseudo-terminal (but behaving ter-
minal), and variously bi- or tripinnately paniculate, with branches ending
in loose to tight corymbs of flowers. The panicles are mostly pyramidal,
but may be reduced to a columnar panicle of corymbs, thus appearing race-
mose in fruit (e.g., Hymenandra sordida). The inflorescence bract is foliose,
membranaceous, nearly sessile and early caducous. The peduncle, primary
rachis and branches are glabrous or furfuraceous lepidote, then glabrescent.
The floral bracts are ovate, lanceolate or narrowly ellpitic, and membrana-
ceous, sparsely to densely furfuraceous at first, then glabrescent. The pedicels
are cylindrical and bear the same vestiture as the inflorescence branch. The
flowers are perfect, bisexual, homomerous, and 5 (—6)-merous, and both perianth
members may be membranaceous, chartaceous or coriaceous, with pellucid
or black punctations and punctate-lineations. The calyx has lobes nearly
free, lanceolate, linear-lanceolate, linear, oblong, or suborbicular, symmetrical
or asymmetrical to slightly auriculate, apically acute, obtuse or rounded,

the margin regular, irregular, or subapically notched, hyaline or scarious,
sparsely glandular-ciliolate, minutely erose or entire (Fig. 1). The corolla is
campanulate, the lobes nearly free, or free, ovate, lanceolate, oblong, or el-
liptic, symmetrical, apically acute or acuminate, erect or reflexed in anthe-

708 Sipa 18(3)

2mm

A B C D

Fic. |. Comparison of calyx lobes size, shape and punctations amoung the Neotropical specie
of Hymenandra. A. ieee oars drawn from M. chinchilla 93. B. Hymenandra empl
drawn from B, sa 13717.C. Hymenandra pittieri, drawn trom K. Sytsma, W. Hahn & T
Antonto 2453. D. Hy ee ilar: iana, drawn from G. Davidse . Herrera 31379. E
ee ee ae from R. Callejas et al, 5683. F. Hymenandra crosby1, drawn fom
L. Andersson & R. Dressler 4279. G. Hymenandra acutissima, drawn from the ho-
lotype, J. Cuatrecasas 17 7 H. Hymenandra sordida, drawn from the holotype, 7. Croat
43538. 1. Hymenandra squamata, drawn from the MO isotype, W. Burger G RB. Stolze 5853.

sis, glabrous, the margin hyaline, and entire. The stamens are connate at
least basally by their filaments to form a staminal tube, and by their an-
thers along the dehiscence sutures. The filaments are connate throughout
their length in the neotropical species, but only basally in the Indo-Malesian
species. The anthers are basifixed, connate along their dehiscence lines, e1-
ther basally at first, or throughout their length, and lanceolate or ovate,
apically long-attenuate to an acute or emarginate tip, or rounded with an
apiculate tip, basally cordate, dehiscent by longitudinal slits, and with connectives
ventrally darkened or prominently black punctate dorsally in che basal portion.
The pistil may be obturbinate or ellipsoid, the style longer than the ovary,

—

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 709

attenuate to a punctiform stigma. The placenta is subglobose to ellipsoid
with relatively few to numerous, pluiseriate ovules in a high anthotactic
spiral. The fruit is a globose to depressed-globose, one-seeded drupe, with
a persistent style, sparsely or densely and prominently, conspicously or in-
conspicuously pellucid or black punctate.
ECOLOGY

Neotropical Hymenandra species are mostly small trees, most commonly found
along ridgetops in moist, wet, cloud or pluvial forests, from near sea level
to 1,400 m elevation. Two species, Hymenandya sordida and H. squamata,
are subshrubs less than one meter tall, but they are ecologically distinct,
the former is known only from cloud forests, and the latter known from
swamp forests. Only one species, Hymenandra stenophylla, is found in low-
land moist forests. Hymenandra wilburiana and H. pittieri are both found in
forests transitional between lowland and premontane wet forests, but the
former species is restricted to ridgetops, while the latter is found along the
forest margin, particularly along watercourses. In the Colombian Chocé,
two species occur that are easily separated ecologically. Hymenandra callejasti
is from the lowland pluvial forest of the northern Antioquian Chocé, while
Hi. acutissima is a riparian species along major rivers along the shores of the
Pacific Ocean. Hymenandra calycosa is known only from premontane wet forests,
from Nicaragua to Panama.

In Mesoamerica, the altitudinal compression or expansion of life zones is
largely a result of exposure to winds, rain and other meteorological events.
Therefore, absolute altitude often has little to do with the life zone present
at a given site. For example, many areas in Nicaragua support vegetation
otherwise found only at much higher altitudes, because there are no taller
mountains around. Fieldwork has shown that even a small mountain of 800
m can support cloud forest. Such appears to be the case of Hymenandra croshyi,
which is restricted to the Caribbean Slope of Panama, at the junction of the
premontane and cloud forest life zones, even though the altitudinal range
of the species is only 100-850 m.

BIOGEOGRAPHY AND IMPLICATIONS FOR THE BOREOTROPICS HYPOTHESIS

Hymenandra, as now defined, is amphipacifically distributed, occurring
from Bangladesh eastward through Malaya to Borneo, and in the Neotropics
from Nicaragua southward through Costa Rica and Panama to the Chocé
of Colombia. A distribution such as this, in particular with the majority of
the species centered in Central America, most closely fits those groups cited
as partial evidence to support what Wendt (1993) discovered in his study
of lowland Mexican wet forests, and what Lavin and Luckow (1993) attrib-

710 Sipa 18(3)

uted to the Boreotropics Hypothesis, proposed by Wolfe (1975) and Tiffney
(1985a, b) to explain the distribution of the “boreotropical flora.” The hy-
pothesis proposes that the biotas of North America and Europe, including
tropical North America, were once more widespread in the northern hemi-
sphere and transgressed the North Atlantic by direct land connections or
over limited water gaps until the late Ecocene or early Oligocene (Lavin &
Luckow 1993). A logical extension to Wolfe’s and Tiffney’s concept, pre-
sented by Wendt (1993) and Lavin and Luckow (1993) is that the boreotropical
flora not only existed in the Eocene, but also left a significant number of
direct descendant lines in present lowland tropical floras of northern Latin
America. Therefore, we would expect that many of the early Tertiary fossil
taxa from both North America and Europe were most closely related to extant
species from tropical southeast Asia, and to some extent, Central America
and the Greater Antilles (Lavin & Luckow 1993). It is the latter notion that
is congruent with the Amphipacific distributional pattern like that found
in our new concept of Hymenandra. This extension to the boreotropics hy-
pothesis was cited by Wendt (1988, 1989, 1993), in discussing the rela-
tionships of Chiangiodendron (Flacourtiaceae), and by Zona (1990) in dis-
cussing the biogeography of Sabal (Arecaceae). A similar distribution for
the genus Adstonia (Apocynaceae) was cited by Gentry (1983), but he did
not invoke the hypothesis per se. Conran (1995), in his study of the Liliiflorae,
found that three taxa defining the Southeast Asian/northern Australasian
clade (Stemonaceae, Hanguanaceae and Uvulariaceae) were widespread northern
taxa that have spread southwards.

The geographic distribution of Hymenandra is entirely consistent with
the area cladogram presented by Lavin and Luckow (1993, Fig. 1), where
South American elements are ultimately derived from among diverse North
American lineages, these lineages having a sister group relationship to
Paleotropical groups. While Wolfe’s hypothesis could be correct even if no
modern descendants of the boreotropical flora were found in the Neotopics
owing to extinction, the three criteria proposed by Lavin and Luckow (1993)
to test the hypothesis were: 1) a center of diversity in North America (in-
cluding “tropical North America” as they define it), 2) an early Tertiary
fossil record in North America, and 3) a pantropical distribution.

For the first test criterion, “North America” includes both tropical and
temperate elements, the tropical ones south of the Tropic of Cancer. In the
case of Hymenandra, it is clear that the majority of the neotropical members
of the genus (H. stenophylla, H. calycosa, H. sordida, H. squamata, Hu ilburiana,
H. crosbyi) are centered in Nicaragua, Costa Rica, or the Caribbean slope of
Panama, with only H. p/ftier? spanning the central montane of Costa Rica
and Panama. Hymenandra callejasii and H. acutissima are restricted to the

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 711

Chocé of Colombia, a floristic province whose northern limit is southeast-
ern Panama. Therefore, the pattern fits the first criterion.

Unfortunately, no data is available for use in the second test criterion
owing to lack of fossils known for the group. As for the last criterion, a
pantropical distribution, the closest related genus to Hymenandva appears
to be the genus Connandrium Mez, known only from New Guinea, on the
basis of possession of anthers fused along the sutures. In connection with
New Guinea and adjacent Australia as significant sources of biota related
to that of North America, Dobson (1996) cites the distribution of the ter-
mite genus Mastotermes and the sister poison frog genera, Pitohui and Phyllobates,
as further evidence, but without citing the boreotropics hypothesis per se. It
is unfortunate that the preliminary cladogram for genera of the family by
Stahl (1996) did not include Hymenandra. The other neotropical genera of
the tribe Ardisieae include Ardisia, pantropically distributed with major centers
of diversity in Malesia and Mesoamerica, Crenardisia Ducke, with species
from northern Mesoamerica and the Amazon Basin, Synardisia (Mast.) Lundell,
a monotypic genus ranging from southern Mexico to Nicaragua, and Gentlea
Lundell, with the majority of species in Mexico and Mesoamerica (only one,
G. venossisima, in the Andes from Venezuela to Peru). Until a phylogenetic
analysis is complete for the tribe, a rigorous test cannot be performed. However,
the pattern of distribution among the genera of the tribe Ardisieae fit the
general pattern one would expect if the distribution was boreotropical

In summary, we hypothesize that Hymenandra appears to fit the overall
pattern consistent with a boreotropical distribution. However, we must emphasize
that until a cladistic analysis among the genera of Myrsinaceae is complete,
there is no reliable way to unequivocally determine if the group is boreotropical
or Gondwanan. However, at this point in our work, we find it useful to
point out the strong correlation and call attention to the value of examin-
ing generic limts on a worldwide basis when preparing treatments for a
large flora such as Flora Mesoamericana.

NOTES ON KEYS, DESCRIPTIONS, AND TERMINOLOGY

The keys are artificial and designed to expedite identification of herbarium
specimens. An attempt has been made to emphasize vegetative characters
to increase the keys’ usefulness with sterile material. The numbers appear-
ing before the taxa refer to their respective position in the key; any correla-
tions with phylogenetic relationships are coincidental. Quantitative and
qualitative data presented in keys and descriptions for floral parts and bracts
were taken from organs rehydrated from herbarium specimens by boiling
in water. Measurements from these range from 10% to 15% greater than
those measurements taken directly from dried material. Data regarding shoot

rise Stipa 18(3)
diameters, inflorescence rachises, pedicels, leaf and fruit shape and size were
taken from dried herbarium specimens.

Morphological terms in this treatment follow Lindley (1848) and Pipoly

(1987, 1992a) for the inflorescence, rachis pedicels and floral parts. Description
of leaf morphology follows fae (1984), trichome description follows Theobald
et al. (1984) and basic cell and tissue terminology follow Metcalfe (1984).

Because of branchlet and leaf dimorphism between shoots in vegetative
vs. reproductive phases, the primary orthotropic axes (stems if a small statured
plant, trunks ifa treelet), are shoots not bearing inflorescences and are termed
“vegetative shoots,” while those plagiotropic branches actually or potentia
bearing flowers or fruit are termed “reproductive shoots”.

_—

ly
Likewise, the leaves
on the trunk or on the branches when they are in a vegetative state, will be
termed, “vegetative shoot leaves” while those leaves on the shoots while they
are in the reproductive phase will be termed, “reproductive shoot leaves.”
All types cited here have been seen by the authors unless noted as “n.v.”

KEY TO GENERA OF MYRSINACEAE IN MESOAMERICA

Calyx closed in bud, it nee ly opening into 2-8 lobes among flowers within
the same inflorescenc

. Calyx open in bud, with (3—)4—5S(—6) lobes, the number of
pc consistent within one anHlones cence:

2. ne and corolla lobes valvate 1 i rufous glandular-
sillac

—

Geissanthus
caylx lobes per

“Blowers bisexual; inflorescences paniculate, longer than the petioles
siete tomentose with stellate or dendroid trichomes; calyx 4
rolla tomentose and glandular-papillose; anthers bright yel
elongate

nd co-
low; style

Parathesis
. Flowers unisexual; inflorescences glomeru

reduced c

—

ate, umbellate or rarely in
orymbs, much shorter than the petioles; shoots glabrous or
when tomentose, then with simple trichomes; calyx and corolla gla-
brous or the corolla sometimes glandular-p

yapillose along the margin
within; anthers cream to white; ope absol ete or reduced dich thesaeeedenneneee Myrsine
Calyx and corolla alone uncial, iml i

so), contorted or ré

NO

ely valvate in Sad: when valvare, the indlonks scence ra-
ae ous ee mers al sillate,

sasally to form a staminal tube adnate to the corolla
tube.
5. Inflorescence lateral (axillary); corolla tube glandular-granulose within
at least at che lesion of the tube and lobes; staminal tube adnate
to corolla at least iu S11 fe Cybianthus
5. Inflorescence terminal or ene functioing as terminal; corolla
tube glandular-puberulent within at base; sta ree f
adnate to corolla at base.

ns free from or barely

6. Anchers connate at least at base; filaments connate in a cube at
least '/2 their length; corolla lobes nearly free Hymenandra

PipoLy AND RICKETSON, Hymenandra and its boreotropical implications 713

6. Anthers free; filaments connate less than 1/5 length; corolla lobes

united up to 3/4 their length Synardisia
. Filaments free from each other and free from or variously adnate to the
se tube, or filaments basally connate to form a staminal tube not
adnate to the corolla tube
7. Filaments adnate to the corolla; anthers less than 3 times longer than
wide
8. Filaments exserted, more than 6 times longer than the anthers;
anthers obcordate; plants of elfin and cloud forests Gentlea
8. Filaments included, shorter than or as long as the anthers; anthers
ovate, lanceolate, elliptic or oblong; plants of montane to low
land forests Stylogyne
7. Filaments free from the corolla; anthers at least 3 times longer than
wide
9. Corolla lobes connate 1/4 their length; ovules uniseriate, the placen-
tation free-centra Ctenardisia
9. Corolla lobes nearly free or connate to 1/5 their length; ovules
pluriseriate, che placentation basal Ardisia

TAXONOMIC TREATMENT

Hymenandra (A. DC.) A. DC. ex Spach, Hist. Veg. P han. 9:374. 1840: A.
DC., Ann. Sci. Nat., Bot., ser. 2, 16:79. 1841.; A. DC. in DC. ’Prode.
8: OL. 1844. Ardisia sectio 2. Hymenandra A, DC., on Sci. Nat. “Bots ser. 2,
2:297. 1834; Trans. Linn. Soc. eee 17:126. 1837. Type: Ardisia Dp nunite Wall.
im Roxb. FI. Ind. Ed. Carey 2:282. 1824, = Hymenandra wallichii A. DC., Ann. Sci.

Nat. Bot., ser. 2, 16:79. 1841.
C p cae Lundell, Wrightia 7:41. 1982. Tyre: Ardisia calycosa Hemsl., Biol. Centr.-

, Bot. 2:292. 1882

Subshrubs to small trees exhibiting Fagerlind’s Architectural Model (Hallé
etal. 1978) up to 8 m tall. Leaves pseudoverticillate, dimorphic; the blades
of ihe orthotropic trunk or stem (1.e., primary axis) which are vegetative
shoots larger with cuneate bases, those of the plagiotropic branches which
are reproductive shoots basally obtuse to somewhat auriculate. Liflorescence
terminal or pseudoterminal, bi- or tripinnately paniculate, pyramidal or columnar,
the ultimate branches bearing flowers in loose to tight corymbs. F/owers
perfect, bisexual, 5(—6)-merous, punctate and/or punctate-lineate; calyx lobes
quincuncial, spreading at anthesis, nearly free, mostly papillose but some-
times hirtellous (Paleotropics) or glabrous (Neotropics), densely and prominently
or conspicuously pellucid or black punctate, the margin glandular-ciliate;
corolla white or pink, the lobes short-connate, quncuncial, lanceolate or oblong
to ovate to narrowly ovate, glabrous or sparsely papillose within toward base,

densely and prominently black punctate or inconspicuously pellucid punc-
tate, the margin entire, erose or erose-fimbriate, sometimes subapically notched,
glandular-ciliate or glabrous; stamens connate, shorter than the corolla lobes,

714 Stipa 18(3)

the filaments monadelphous, fused basally 2/3 their length and reunited
apically, fused basally but with apical portions free, or fused along entire
length, adnate basally to the corolla tube, the anthers connate basally for 1/
4—2/3, or along entire length along the thecal margins, narrowly oblong,
linear-lanceolate or lanceolate, the connectives darkened or prominently black
punctate abaxially; piscil obturbinate, the ovary ovoid, the style slender,
elongate, the stigma punctiform; placenta subglobose, S—24 ovules, at times
appearing uniseriate (but in a high anthotactic spiral), or clearly pluriseriate.
Fruit 1-seeded, globose or depressed-globose, the exocarp densely and prominently
or conspicuously punctate, the style base persistent.

Distribution.—A genus of 16 species, from Bangladesh eastward to Myanmar,

south to Malaya and Borneo in the Paleotropics; Nicaragua southward through
Costa Rica and Panama to the Chocé of Colombia in the Neotropics, from
near sea level—1,400 m elevation.
Members of Hymenandra are subshrubs to small trees in the lowest
level arte forest understory, from the lowland to premontane transition to
cloud forest, frequently on ridge tops or steep slopes, and near small water-
courses. They are at best locally common {see ECOLOGY section}.

Etymology.—Hymenandra is a Greek compound, derived from the words
Hymen, or membrane, and andros, of the anthers, to describe the membranaceous
connective tissue joining the anthers at their dehiscence suture.

KEY TO NEOTROPICAL SPECIES OF HYMENANDRA

Calyx lobes oblong or ie oblong to narrowly lanceolate, at least 3—5
times — than wide at maturity.
2. Calyx lobes shorter than the corolla lobes, 2.3-4.7 mm long, 0.9-1.1 mm

on

wide, the cag erose, hyaline, densely glandular-ciliolate; corolla lobes
5 n long 1. Hymenandra enash

2. Calyx ee longer than the corolla lobes, 6-9 mm long, (1.6—)2—

mm wide, the margins entire, glabrous; corolla lobes 7-8.2 mm lon
Hymenandra calycosa

Calyx lobes ovate or suborbicular, usually less than 2 times longer than wide
at maturity.
3. Inflorescence a columnar panicle of corymbs, often appearing racemose i
3.H

ruit : eae sordida
Dy Inflorescence a pyr amidal icle of b ippeat ing g paniculate in fruit.
4. Calyx lobes asymmetric, notched subapically:
>: Sa .2—1 m tall; abaxial leaf surface densely and Bea en
furfuraceous ‘ pidote, appearing velutinous; stamens 2.6—3 mm | one
the anthers apically rounded, apiculate at tip, 1.6-1.8 mm a 0.8
0.9 mm wide Hymenandra squamata

. Trees, 1-8 m call; abaxial leaf surface glabrous to sparsely and mi-

MN

nutely nea lepidote, not appearing veluctinous; stamens 6—

9.5 mm long, the anthers apically long-attenuate to an acute or emarginate
1.4—2 Wi

tip, 5.5—-8 mm long, 2mm

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 715

6. Calyx lobes 1.4—-1.8 mm long; stamens 6—6.5 mm long; anthers
He mm long; vegetative shoot leaf blades 5.5—22.2 cm long,
78 cm wide, reproductive shoot leaf blades 7.2-17.3 cm long,
ae cm wide 5. Hymenandra aes
6. Calyx lobes 2.2—2.8 mm long; stamens 8.7—9.5 mm oe anther
u oe 8 mm long; vegetative shoot leaf blades 33.7—72.2 mon
—16.5 cm wide; reproductive shoot leaf blades 3.9—34.3
ae 1.2-11.8 cm wide 6. cance pittieri
4. Calyx lobes symmetric, not subapically notched.
7. os or flowering shoots 7-10 mm in diam.; petioles (0.6—)1—
cm long; calyx lobes 2—2.8 mm long................ 7. Hymenandra callejasii
re oo or flowering shoots 1.5—5.5 mm in diam.; petioles 1.6—
3.9 mm long; calyx lobes 0.9-1.6 mm lon
8. Calyx lobes 1.4—1.6 mm long; rete terminal, 9-16.5 cm
long, secondary branches 5—5.5 cm long; fruit 8-12 mm in diam.
menandra acutissima
2

8. Calyx lobes 0.9-1.2 mm long; inflorescences subterminal,
8.8 cm long, secondary branches 1.3—3.5 cm long; fruit 4.5—6.5
mm in diam. 9. Hymenandra crosbyi

i a stenophylla (Donn. Sm.) Pipoly & Ricketson, comb. nov.
(Figs Ha Donn. Sm., Bot. Gaz. 24:395. are ies
oe pee Ral Phytolosia 49:351. 1981. Type. COSTA RICA. Lr
ests of Shirores, Talamanca, 300 ft. [91 m}, Feb 1895 (A), H. Prttier G i 2
9173 (HoLoTYPE: US (LL Neg. # 71-109, US Neg. # 2388); isorypes: CR n.v., US).

Ardisia oblanceolata Standl., Publ. Field Colombian Mus., Bot. Ser. 4:249. 1929, syn.

ov. Type. PANAMA. Bocas Det Toro: Region of iene: without elev., Jan—

ae 1928 (fr), G. Cooper 370 (HoLoTyPE: F (F Neg. # 68229, LL Neg. # 1971-75);
ISOTYPES: G (fragment), K, NY, US (US Neg. # 2378)).

Tree 2—6.1 m tall, to 7.5 cm in diam. Trunk and vegetative shoots terete, 7—
9.5 mm in diam., minutely scattered furfuraceous lepidote, early glabres-
cent; reproductive oe similar, but 1.5—4 mm in diam. Vegetative shoot leaf
blades chartaceous, oblong to elliptic, 21.2—30.5 cm long, 3.5—8.5 cm wide,
apically long-acuminate, the acumen 1—1.9 cm long, gradually tapering to
a cuneate base decurrent to base of petiole, midrib slightly raised above,
prominently raised below, secondary veins 22—32 pairs, slightly depressed
or not visible above, slightly raised below, prominently punctate, scattered
minutely furfuraceous lepidote, early glabrescent, the margins entire, flat;
petioles marginate, 0.5—1.5 cm long, minutely scattered furfuraceous lepi-
dote, early glabrescent; reproductive shoot leaf blades similar to the vegetative
ones but 6.1—22.5 cm long, 2.1—5.2 cm wide, secondary veins 7—28 pairs;
petioles similar to vegetative ones but 0.5—2.2 cm long. Inflorescence termi-
nal, pinnately or bipinnately paniculate 3-10 (-15.5) cm long, pyramidal,
secondary branches 0.8—3(—6.2) cm long, minutely scattered furfuraceous
lepidote, early glabrescent, the branches ending in 4—8-flowered corymbs;
inflorescence bracts early caducous, membranaceous, oblong, 5.6—8.8 mm

716 Stipa 18(3)

Fic. 2
Detail of abaxial leaf surface. ail
stamens, sho

2. Hymenandra stenophylla ees Sm.) Pipoly & Ricketson. A. Reproductive shoot. B.

of inflorescence. D. Detail of flower. E. Detail of

wing abaxial (Gekiand ea (right) surfaces. F Ovules. G. Fruit. A~B drawn

from the holotype, H. Prttier & A. Tonduz 9173.C drawn from L.D. Gomez et al. 20415. D-
F drawn from B. Hammel 13717. G drawn from J. Gomez- ae ito 9538

Qu

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 717

long, 0.6-1.2 mm wide, apically acute to rounded, midrib slightly raised
above and below, secondary veins not visible, prominently punctate, scat-
tered minutely furfuraceous lepidote, the margin entire, flat; floral bracts
0.5-2.2 mm long, 0.3—1.1 mm wide, otherwise as the inflorescence bracts;
pedicels cylindrical, 0.6-1.2(-1.6) mm long, scattered minutely furfura-
ceous lepidote, early glabrescent. F/owers S-merous, white or rarely pink;
calyx lobes almost free, chartaceous, narrowly lanceolate to narrowly ob-
long, 2.3-4.7 mm long, 0.9-1.1 mm wide, symmetrical, apically very nar-
rowly acute to subulate, densely and prominently black punctate and punctate-
lineate, glabrous, the margin irregular, erose, hyaline, sparsely glandular-ciliolate;
corolla lobes membranaceous, 5—6.5 mm long, the tube 0.2—0.5 mm long,
the lobes lanceolate 4.5—6.3 mm long, 2—2.8 mm wide at base, apically
acute, reflexed in anthesis, prominently pellucid punctate, the margins hyaline,
entire, glabrous; stamens 3.5—5 mm long; filaments connate throughout
their length into a chartaceous elobate tube, the tube 0.3—0.8 mm long,
adnate to the corolla basally, epunctate, glabrous; anthers connate by their
sutures, at least at first, lanceolate, 3—-4.5 mm long, 0.7—1.1 mm wide at
base, sessile on staminal tube, apically apiculate, basally cordate, longitu-
dinally dehiscent by narrow, sublatrose slits, the connective densely and
prominently black punctate; pistil obturbinate 5.3—7 mm long; ovary 1.2—
1.5 mm long, glabrous; style 3.8—4.4(-5.8) mm long, erect, inconspicu-
ously pellucid punctate, glabrous; stigma punctiform; placenta ellipsoid,
0.4-0.7 mm long, 0.2—0.5 mm in diam., apically apiculate; yes 13-17,
pluriseriate. Frait obovoid to globose, 4.8-5.8 mm long, 3.8—4.5 mm in
diam., densely and prominently punctate, the style base persistent.

Damabaiia —Limén, Costa Rica and Bocas del Toro, Panama, from 25—
500 m elevation.

Ecology and conservation status. —Known only from primary lowland tropical
moist forests, collected from the understory along ridgetops, Hymenandra
stenophylla should be considered threatened.

Etymology.—The epithet ‘stenophylla’ comes from the Greek “sfevos” meaning
narrow, and ‘phy//us’ referring to the leaves. It describes the very narrow
leaf blades of the taxon.

ae

Specimens examined. COSTA RICA. Limén: panna Valley, without elev., 1927
(ster.), G. Cooper 12 (US); 7 km SW of Bribrf, 100-250 m, 4 May 1983 (fl), L. Gomez et al.
20415 (LL, MO); Sitio de exploraci6n petrolera, Suretka, Talamanca, 250 m, 6 Jul 1983 (fr),
J. Gomez-Laurito 9538 (F); Headwaters of Quebrada Mata de Limon, central fork, and hills
between central and W forks, Finca Anai, Sixaola region, 09° 35'N, 82° 39! W, 25-40 m,
19 Nov 1984 (fr), M. Grayum et al. ae (MO, NY); sae Cerere Reserve and vicinity in

Valle la ey of Finca Conce n, from station to top of ridge Miramaror Los Jabillos,
09° 42'N, 83° 02' W, 140-500 m, a Au 1985 (fr), B. Pome Grayum 14342 (MO);
Reserve Biol. ais toy-Cerere, near the on, Valle de la Estrella, 09° 40' 30" N, 83° O1'

30" W, 100 m, 20 Sep 1991 (fr), A. ee 7 (MO); Forets de Tsaki, Talamanca, 200 m,

718 Sipa 18(3)

Apr 1895 (fl) A. Tonduz 9586 (G 3-sheets, US). PANAMA. Bocas del Toro: Along oil pipe-
line road between continental divide and Chiriqui Grande, second bridge over Rio Guabo
along dirt acces{s] road, 08° 50—55' N, 82° 9-15! W, 120-200 m, 1-2 May 1985 (fl), B.
Hammel 13717 (MO), Filo Almirante along trail to Risc6 Abajo, 3 km SW of town of Almirante,
100-200 m, 3 Jan 1975 (fr), M. Nee & B. Hansen 14099 (MO); Vicinity San San River,
Davao Farm, United Fruit Co., Almirante, 26 Aug 1940 (fr), R. Sezbert 1572 (MO, US).

Hymenandra stenophylla appears to be most closely related to H. calycosa
by virtue of its oblong or narrowly oblong to narrowly lanceolate calyx lobes.
However, the erose and sparsely glandular-ciliate calyx lobe margins, the
calyx lobes smaller than the corolla lobes, and generally smaller flowers,
easily separate the two species.

Populations corresponding to the type of Ardisia oblanceolata are notable
only for the slightly larger and more prominient punctations on the leaves
and slightly shorter pedicels. It is otherwise identical to Hymenandya stenophylla.

2. Hymenandra calycosa (Hemsl.) Pipoly & an comb. nov. (Fig
LA, 3). Ardisia me Hemsl., Biol. Centr.-Amer 292. 1882. Chontalesia
ie (Hemsl.) Lundell, Wrightia 7:42. 1982. Typ PE. a GUA. CHONTALES:
Chontales, a au. 1867-1868 (fl), R. Tate 239 {454} (HoLoryer: K; 1isorypes:

Shrub or tree 1-4 m tall. Trunk and vegetative shoots terete, 4—7 mm in diam.,
minutely and densely furfuraceous lepidote at first, early glabrescent; repro-
ductive shoots as in vegetative ones but |—4 mm in diam., minutely scattered
furfuraceous lepidote. Vegetative shoot leaf blades coriaceous, narrowly ellip-
tic, 7.5—20.3 cm long, 2.8—5.2 cm wide, apically long-acuminate, the acu-
men 0.5—2.2 cm long, gradually tapering to the cuneate base, decurrent to
petiole base, midrib slightly raised above, prominently raised below, sec-
ondary veins 7—21 pairs, slightly depressed or not visible above, prominently
raised below, prominently punctate, glabrous above, minutely scattered
furfuraceous lepidote below, early glabrescent, the margins crenate; peti-
oles marginate, 0.2—0.8 cm long, glabrous above, minutely scattered fur-
furaceous lepidote below, early glabrescent; reproductive shoot leaf blades as in
vegetative ones but 7.2—15.7 cm long, 1.6—4.1 cm wide, secondary veins
7—20 pairs; petioles like the vegetative ones but 0.3—0.9 cm long. Inflorescence
terminal, pinnately or bipinnately paniculate, 4.5—14 cm long, pyramidal,
secondary branches 2.7—4.3 cm long, minutely scattered furfuraceous lepi-
dote, early glabrescent, terminating in 3—7-flowered corymbs; inflorescence
bracts early caducous, membranaceous, oblong, 5.5—9 mm long, 0.7—1.1
mm wide, apically acute, basally sessile, midrib slightly raised above and
below, secondary veins not visible, prominently punctate, mostly glabrous
above, densely minutely furfuraceous lepidote below, the margins entire,
flat; floral bracts similar to inflorescence bracts but lanceolate to narrowly
elliptic 2.8—4.1 mm long, 0.4—1.2 mm wide; pedicels cylindrical, 7-15

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications rie

Pic. 3. Hymenandra calycosa (Hems!l.) Pipoly & Ricketson. A. Reproductive shoot. B. De-
tail of abaxial leaf surface. C. Detail of inflorescence. D. Detail of flower. E. Detail of sta-
mens, showing abaxial (left) and adaxial (right) surfaces. E ovules. G. Fruit. A-B drawn
from BM isolectotype, R. Tate 239 {454}. C-F drawn from M. Chinchilla 93. G drawn
from R. Villalobos 37.

720 Sipa 183)

mm long, minutely scattered furfuraceous lepidote, early glabrescent. Flowers
5-merous, 6.9-8 mm long; calyx lobes almost free, coriaceous, narrowly
lanceolate to lorate, 4.2—9.7 mm long, 1.2—2.1 mm wide, symmetrical, apically
acute to rounded, prominently punctate, appearing prominently because
of the prominently brown punctations, scattered lepidote outside at the base,
the margins entire; corolla lobes basally connate, membranaceous, the tube
1.5—1.8 mm long, elliptic, the lobes 5.4—6.2 mm long, 2.2—2.4 mm wide
at the middle, apically acute to acuminate, reflexed in anthesis, prominently
pellucide punctate, the margins hyaline, entire, glabrous; stamens 4.8—5.1
mm long; filaments connate into a chartaceous elobate tube, the tube 0. 4—
0.5 mm long, adnate to the corolla basally, epunctate, glabrous; anthers connate
by their sutures, at least at first, lanceolate, 4.5—4.7 mm long, 1.2—1.5 mm
wide at the base, sessile on the staminal tube, apically apiculate, basally
cordate, longitudinally dehiscent by narrow, sublatrose slit, the connective
conspicuously raised, pellucid punctate; pistil obturbinate 6.5—7.2 mm long;
ovary 1.2—-1.4 mm long, glabrous; style 5.3—5.8 mm long, erect, epunctate,
glabrous; stigma punctiform; placenta ellipsoid, 0.6—-0.8 mm in diam., apically
apiculate; ovules 11-19, pluriseriate. Frat elliptic, 6.2-7 mm long, 9.2—
5.8 mm in diam., nearly epunctate, the style base persistent.

Distribution. —Jinotega, Chontales, Zelaya and Rio San Juan, Nicaragua
and Alajuela and Guanacaste, Costa Rica. It grows from (10—)450-1,100
m elevation.

Ecology and conservation status. —This species normally occurs in premontane
wet forests, especially along ridgetops. The locations where the species has
been found are actually premontane even though their absolute elevations
are at times quite low. This phenomenon is due to the relative flatness of the
Atlantic Slope in Nicaragua, where even elfin forest has been found under 1,000
m elevation. With increasing habitat destruction, and its consequent desertification,
Hymenandra calycosa should be considered a threatened species.

Etymology.—The specific epithet refers to the calyx, which is always longer
than the corolla and is accrescent in fruit.

Specimens examined. NICARAGUA. Chontales: Cerro Oluma, on poe top of Cor-
dillera Amerisque, 750 m, 4 Jan | 984 (fr), A. a ét al, 43939 (DUKE, NY); Cerro
Oluma, ca 3 3 km SW de Ciudad ane 12° 18'N, 85° 20' W, 700- ee a 1984 (fl),
A. Grijalva et al. 3393 (M); Cerro El Portillo, ca. 3 km al NE de Ciudad Cuapa, 758 m, 1
Mar 1984 (fl, fr), A. eae | D, las 3652 (MO). Jinotega: Kilambé, Cerro San Pedro,
13° 36' N, 85° 39! W, ca. 600-800 m, 25 Mar 1981 (fl), P. : oreno 7532 (MO, NY); Las
Brisas, Comarca ‘eilambe. 13° 351.36! N, 85° 39' O, 930 m, 13 Jul 1980 (A), - spa
159 (MO, NY). Rio San Juan: Boc: a de Sdbalo, trail to Buena i 11°03'N,8
ca. 70 m, 24 Mar 1985 (A, fr), 2. Moreno 25630 (MO); Sdbalo, 1 km al N de Rio - ae
11° 02' N, 84° 27! W, ca. 100 m, 9-10 Jul 1985 (fr), P. Moreno 26053 (MO); Bocas de
Sdbalo, 11° 03' N, 84° 27 W, 0 0-100 m, 14 Mar 1987 (ster.), PB. Moreno 26753 (MO).
Zelaya: Rio Punta Gorda, Atlanta, desembocadura del Cafio el Guineo, 11° 33' N, 84°

=

PipOLy AND RICKETSON, Hymenandra and its boreotropical implications 72

02' W, ca. 10 m, 11 Nov 1981 (fr), P Moreno & J. Sandino 12853 (MO). COSTA RICA.
Alajuela: Ca. 3 km NNE of Bijagua along the new road to Upala, 10° 45' N, 85° 03' W,
450 m, 7-8 Nov 1975 (fr), W Burger & R. Baker 9869 (F); E slopes of Volcdn Miravilles,
W of Bijagua, near the Rio Zapote, 10° 44' N, 85° 05' W, ca. G00 m, 11-12 Feb 1982 (fl),
W. Burger et al. 11687 (B, LL, NY); Cataratas (Los Angeles) de San Ramon, without elev.,
17 Apr 1925 (fl), A. Brenes 21243 (NY); San Carlos Cant6n, Betania de Cutris, 150 m, 26
Feb 1983 (fl), A. Carvajal U. 392 (DUKE, LL, MO, NY); El Silencio de Tilaran, without
elev., 31 May 1932 (A, fr), A. Brenes 15655 (F, NY); Upala Canton, P.N. Guanacaste, Cor-
dillera de Guanacaste, Estacién San Ramon, Dos Rios, 10° 52'50" N, 85° 24' 05" W, 550
m, 3 Apr 1995 (fl), M. Chinchilla 93 (MO), 100 (MO); Along road between Canas & Upala
4 km NNE of Bijagua on slopes leading into Rio Zapote, ca. 400 m, 24 Jun 1976 (A), T.
Croat 36246 (LL, MO, NY); Upala Cantén, P.N. Guanacaste, Cordillera de Guanacaste,
Estacién San Ramon, bosque en el sendero El Argentino, 10° 52' 40" N, 85° 24! 15" W,
550 m, 22 Mar 1994 (fl), D. Garcfa 312 (MO); Santa Marfa National Park, road down
Caribbean slope | km E of summit of road, 4 km W of E side of park, 4 km E of colored
house at junction of road to Hacienda Santa Marfa, 10° 37! N, 85° 17! W, ca. 600 m, 7 Feb
1978 (fl), R. Liesner 5066 (LL, MO, NY); 2 km N of Santa Rose, 15 km N of Boca Arenal
on Quesada-Muelle San Carls-Los Chiles road, 10° 38' N, 84° 31' W, ca. 100 m, 28 Apr
on (fl), R. Liesner et al. 15041 (MO); Upala Cantén, P.N. Craik Cordillera de Giane iste,
Estacién San Ramé6n, sendero El Argentino, camino al Volcan Cacao, 10° 53' 50" N, 85°
24' 29" W, 850 m, 27 Feb 1996 (Al), J. Morales 5305 (MO); Moravia de Cutris-San Carlos,
without elev., without date (ster.), L. Poredat et al. 3681 (F); Cantén San wan Cuenca de
pe ae 20 km después de Pavén, Finca Elky Marfa, 10° 52' 30" N, 84° 30! 25
W, 10 , 16 Mar 1997 (fl), A. Rodriguez & N. Zamora 2067 (CR, iE ie MO): Villa
eee 850 m, 21 Feb 1939 (fl), A. Smith 1630 (F, GH, MICH, NY), 1726 (F, MICH),
2609 (F, MO); On the slope 2 Volcan Rincon de la Vieja, along the road N from Liberia,
ca. 5 km from the park, 400-700 m, 1 Mar 1985 (Al), C. Taylor 4551 (DUKE, LL- oe of
DUKE); 2 km N of Bijagua a7 coals: de and on trail to San Miguel or 7 km i
Naranjo, 400-500 m, 8 Jul 1976 (fl), J. Utley & K. Utley 5318 (DUKE). eee ae
order: Border region on the slopes of Volcan Miravalles, along road N of Bagaces, from
Guayabo to Aguas Claras, 400-600 m, 2 Mar 1985 (A, fr), C. Taylor & C. ‘Shoat 4630
(DUKE). Guanacaste: La Cruz Cantén, Parque Nacional Guanacaste, Cordill
Escacién Pitilla, Sedero los Memos, 10° 59! 26" N, 85° 25! 40" W, 740 m, 15 a 1995 aL
fr), E. Alfaro 169 (MO); Liberia Cantén, Ps Nason Guanacaste, Cordillera de Guanacaste
Nueva Zelandia, Estaci6n San Ramén, 10° 52'50" N, 85° 24' 05" W, 550 m m, 23 Mar 1994
(fl, fr), D. Garefa 112 (MO); Tilacsee tidge above town ca. 2.5 mi, without elev., 6 Oct
1971 (fr), A. Gentry 2011 (LL, MO); Ridge to N of Rio Las Flores, ca. 1 km E of Rio Tenorio,
Hacienda Montezuma, 10° 40.5'N, 85° 04.5' W, 475-500 m, 24 Jan 1985 (fr), M. Grayum
& G. Herrera 4852 (MO); Forest along Rio San Juan, lower W Pacific slopes of Volcdn Tenorio,
nda Montezuma, 10° 40.5' N, 85° 03' W, 600-700 m, 25 Jan 1985 (fl), M. Grayum
et al. 4969 (LL, MO); El Dos de Tilardn, 4 km N, Cerro La Chirripa, Atlantic slope, 10°
25' N, 84° 50' W, 1,000 m, 12 Apr 1986 (A, fr), W. Haber et al. 4430 (MO), 4432 (LL),
4447 (MO); La ane ridge, 4 km NE El Dos de Tilaran, continental divide, 10° 25'N,
84° 53' W, 1,000 8 May | ee (fl), We Haber et al. 4752 (MO), 4759 (MO), 4848 (MO):
Tilaran, Rio Chigien Quebrada Gr: Zona Monteverde Finca Lopez, Vertiente Atlantica,
10° 25'N ° 53' W, 700-— ae m, 6 Jun 1987 (fl), W Haber & E. Bello C. 7433 (MO);
Pena ee Guanacaste, Estacién Pitilla, camino al E de la Estacion, 11° 02' N, 85°
25.3' W, 600 m, 24 May 1 989 (fl, fr), B. Hammel et al. 17370 (BRIT, MO); Santa Cecilia,
La Cruz, Estacién pitilla, Faldas Cerro Orosilito, 10° 57' N, 83° 28' W, 700 m, 14 Jun

522 Sipa 18(3)

1989 (fl), Q. Jiménez et al. 709 (BRIT, MO); Santa Elena to Tilardn road, ca. 24 km from
Santa Elena, ca. 1,000 m, 17 Feb 1984 (A, fr) Khan et al. 1141 (BM 2-sheets); Liberia See
Parque 7 Guanacaste, Cordillera de Guanacaste, Estaci6n Cacao, El Cafon, 10° 5
43" N, 85° 28' LO" W, 1,100 m, 10 Feb 1995 (fr), M. Lobo 47 (MO); Estacion pitilla, .
Cruz, eee Nacional Guanacaste, 10° 59' 26" N, 85° 25! 40" W, 700—1,000 m, 7 Nov

1990 (fr), C. Got 214 (MO); Liberia Cantén, Parque Nacional Guanacaste, Cordillera
de Guanacaste, Nueva Zelandia, Estaci6n San Ramé6n, sendero al Volcan Cacao, 10° 53'
50" N, 85° a 29" W 850-950 m, 23 Mar 1994 (fl), J. Morales et al. 2544 (MO), Forest at
top of divide 2 4km NW of Tilardn, without elev., 27 Feb 1972 (fl), P. Opler 580 (F); Liberia
Cantén, Parque Nacional Guanacaste, Cordillera de Guanacaste, Estaci6n Cacao, Sendero
Nayo, 10° 55' 43" N, 85° 28' 10" W, 1,000 m, 9 Feb 1995 (fl), A. Picado et al. 51 (MO),
Hacienda Tenorio, without elev., 17 Feb 1956 (A), B. Schubert 1079 (A, F); El Silencio, near
Tilardn, ca. 450 13 Jan 1926 (fl), P. Standley & J. Valerio 45539 (US), 45545 (US); La
Tejona, N of T ee 600-700 m, 25 Jan 1926 (fr), P. Standley & J. Valerio 45862 (US);
Quebrada Serena, SE of Tilaran, ca. 700 m, 27 Jan 1926 (fl), P. Standley & J. Valerio 46181
(US), 46221 (US); El Silencio, 800 m, 15 Aug 1923 (fr),_/. boned sais hae Liberia Cantén,
Parque Nacional Guanacaste, Cordillera de Guanacaste, Est Cacao, Cerro Cacao, 10°
55' 43" N, 85° 28' 10" W, 1,100 m, 8 Feb 1995 (A, fr), R. Villalobos = (MO

—_—

Hymenandra calycosa is closely related to H. stenophylla, but 1s very easily
recognized by the calyx lobes entire, longer than the corolla and accrescent
in fruit, larger corollas and crenate leaf margins.

33 rs nae sordida (Lundell) Pipoly & ee comb. nov. (Figs.

1H, 4). Auriculardisia sordida Lundell, Wrightia 7:27 2 lida (Lundell)

Lundell, Phytologia 61:66. 1986, nomen. invalidum. “Andi snide (Lundell) J.B.

— eae ae 111. 1997 {1998]. Type. COSTA RICA. ALajueLa: Finca

Los Ensayos, ca. 11 W of Zarcero, ca. 850 m, 15 fan 977 (Al), T. Croat 43538
one yPE: LL; IsoTyPEs: n.v., MO

syle 0.4-1 m tall. Vegetative shoots unknown, reproductive shoots ter-
ete, 4-7 mm in diam., densely appressed furfuraceous lepidote. Vegetative
ae leaves unknown; erage shoot leaf blades chartaceous, elliptic to
oblanceolate, 16—29.5 cm long, 4.5—-10.8 cm wide, apically acuminate, the
acumen 0.5—1.8 cm long, basally atcenuate, the midrib depressed above,
prominently raised below, the secondary veins 35—70 pairs, slightly depressed
or not visible above, prominently raised below, abaxially densely appressed
furfuraceous lepidote along and near the midrib, adaxially glabrous, the margin
entire, inrolled; petioles marginate, 0.5—2.1 cm long, glabrous above, densely
appressed furfuraceous lepidote below. Inflorescence terminal, pinnately or
bipinnately paniculate, 4.2—10.5 cm long, a columnar panicle of 7—14-flowered
corymbs, appearing racemose in fruit, secondary branches 0.5—2.3 cm long,
the rachis densely appressed furfuraceous lepidote, glabrescent; inflorescence
bracts early caducous, membranaceous, lanceolate, 1.4—2.8 cm long, 2.2—
m wide, sessile, midrib slightly raised above and below, secondary

veins obscure to not visable above and below, scattered furfuraceous lepi-
dote to mostly glabrous below, densely appressed furfuraceous lepidote below,

PIPOLY AND RIcKETSON, Hymenandra and its boreotropical implications 723

Fic. 4. Hymenandra sordida (Lundell) Pipoly & Ricketson. A. Reproductive shoot. B. Detail
of abaxial leaf surface. C. Lepidote scale from above and from the side. D. Detail of inflorescence.
E. Detail of flower. FE. Detail of stamens, showing abaxial (left) and adaxial (right) surfaces.
G. ovules. H. Fruit. A-D & G drawn from holotype, 7. Croat 43538. E-F drawn from B.
Hammel et al. 14078. H drawn from A. Molina R. et al. 17525.

724 Sipa 18(3)

the margins entire, ciliolate; floral bracts 1.1—1.7 mm long, 0.3-0.5 mm
wide, otherwise as in the inflorescence bracts; pedicels cylindrical, 5.5—8.7
mm long, scattered to densely appressed furfuraceous lepidote, glabrescent
with age, persistent. F/owers pendent, 5-merous, reddish-violet; calyx lobes
almost free, chartaceous, suborbicular, 1.9-2.2 mm long, the lobes 1.2—

1.5 mm long, 1-1.2 mm wide, asymmetric, apically rounded, subapically
notched, conspicuously black punctate medially, sparsely furfuraceous lepidote,
glabrescent, the margin irregular, somewhat erose, hyaline, densely glan-
dular-cilioate, glabrescent; corolla lobes connate basally, membranaceous,
lanceolate, 3.2—3.7 mm long, the lobes 2.3—2.5 mm long, 1.2—1.3 mm wide
near the base, apically acute, reflexed in anthesis, conspicuously, black- punctate,
the margin erose, hyaline, entire, glabrous; stamens 2.4—2.5 mm long; fila-
ments connate into a chartaceous elobate tube, the tube 1.0—1.1 mm long,
adnate to the corolla basally, epunctate, glabrous; anthers connate by their
sutures, at least at first, ovate, 1.4—1.7 mm long, 0.7—0.9 mm wide at base,
sessile on the staminal tube, apically apiculate, basally cordate, longitud-
nally dehiscent by narrow, sublatrose slits, the connective conspicuously reddish
punctate; pistil obturbinate, 2.8—3.8 mm long; ovary 0.5—0.6 mm long,
glabrous; style 2.3—3.2 mm long, erect, inconspicuously black punctate,
glabrous; stigma punctiform; placenta ellipsoid, 0.1—0.3 mm long, 0.1—
0.15 mm in diam., apically apiculate; ovules 6-9, pluriseriate. Frast de-
pressed-globose, 6—6.5 mm long, 6.5—7.5 mm in diam, inconspicuously
black punctate, the style base persistent.

Distribution —Alajuela and Guanacaste Provinces, Costa Rica, from 500—
1,200 m elevation.

Ecology and conservation status —Hymenandra sordida is a ridgetop species
known only from primary and mildly disturbed cloud forests, and should
thus be considered threatened.

Etymology.—The epithet comes from the Latin word ‘sordidus’, meaning
nel soiled or a

COSTA RICA. Alajuela: Reserva Biol6gica Monteverde Rio Penas
Bienen. Laguna Poco Sol, 10° 21' N, 84° 40' W, 500-800 m, 6 Aug 1989 (Al), E. Bello C.
1059 (MO); San Carlos, San Miguel de San Isidro, 600 m, 29 Jun 1985 (fl), W. Haber G& E.
sae 1714 MO), Pefias Blancas river valley NE of San Carlos, along road from Jabillos
22'N, 84° 38' W, 650 m, 29 Jun 1985 (fl), B. Hammel et al, 14078 (MO); Canton i.
Ups Bijagua, Finca Montezuma, Ladera Sur de Cerro Montezuma, 10° 41' 10" N, 85°
40" W, 600 m, 18 Jul 1993 (fl), G. Herrera 6279 (F, K, MO); 15 km NW of Arenal by
air, 2 km NW of Nuevo Arenal on road to Tilaran, oy n3 a NE on nie to San Rafael de
Guatuso, then 2 km W on road to Finca Cote, 10° 34.5' N, 84° 54' W, 700 m, 30 Apr
1983 (A), R. Liesner et al. 15108 (MO); Near La ean a 8kS ot a a ous ada, 1,200 m,
19 Feb 1966 (fr), A. Molina R. et al. 17525 (FB). eel oe eee Guanacaste,
La Cruz, 9 km al S de Santa Cecilia Estacion Pitilla, 10° 59! 26" N, 85° 25' 40" W, 700 m,
13 Sep 1990 (fl), C. Chituen 71 (MO); Parque Nac ronal Guanacaste, a i6n Pitilla y Sendero

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 725

El Mismo, 10° 59' 26" N, 85° 25! 40" W, 700 m, 15 Jun 1991 (A, fr), B Rvos 374 (INB,
MO); Parque Nacional Guanacaste, Cordillera de Guanacaste, Estacién Pitilla, Santa Cecilia,
9 km S, 10° 59' 00" N, 85° 25' 40" W, 700 m, 20 Aug 1993 (fl), V. Ramirez et al. 62
(BRIT, CR, INB, MO); N slopes of the Volcdn Orosf along lower reaches of broad ridge
running to the N, 650 m, 18 Jan 1968 (fr), R. Wilbur & D. Stone 9745 (LL).

Hymenandra sordida appears to be most closely related to H. squamata but
may be immediately distinguished by the columnar shape of the panicle,
and the larger sepals with merely erose (not erose-fimbriate) margins.

4. Hymenandra squamata (Lundell) Pipoly & Ricketson, comb. nov. (Figs.
LI, 5). Avriculardisia squamata Lundell, Phytologia 56:19. 1984, Ardisia Squamata
Gindelivtas dell, Phytologia 61:67. 1986, nomen. invalidum. isd all (Lundell)
J.F. Morales, Phytologia 83:111. 1997 [1998]. Typr. COSTA RICA. Herepia: Original
forest near the Rfo Puerto Viejo, ca. 2 km upstream from the confluence with Rio
Sarapiqui, formerly “Finca La Selva” of L.R. Holdridge, 10° 26' N, 84° 00' W, 100
m, 14— an i (fA), W. Burger & R. Stolze 5853 (HOLOTYPE: F; isotypes: CR n.v.,
DUKE, 5, L O).

Subshrubs 0.2-1 m tall. Trunk and vegetative shoots terete, 3—5 mm in diam.,
somal appressed furfuraceous lepidote, glabrescent; reproductive shoots ies
2.5 mm diam., densely and minutely appressed furfuraceous lepidote, per-
sistent. ee leaf blades membranaceous, elliptic, 7.4—22.4 cm long,
(1.2—)3.5-6.9 cm wide, apically short to long-acuminate, the acumen ().9—
1.9 cm long, basally attenuate, midrib depressed above, prominately raised
below, secondary veins 18—26 pairs, slightly depressed or not visible above,
prominently raised below, densely appressed furfuraceous lepidote along and
near the midrib below, appearing velutinous, glabrous above, the margin
entire, inrolled; petioles marginate, 0.4—1.1 cm long, glabrous above, densely
appressed furfuraceous lepidote below; reproductive shoot leaf blades similar
to the vegetative ones but (3—)9.0-13.5 cm long, (0.9—)2.2—3.5 cm wide,
tha acumen 0.6—1.1 cm long, basally obtuse; petioles like those of the veg-
etative but 0.2—0.6 cm long. Inflorescence terminal, pinnately or bipinnately
paniculate, 1.4—5.2 cm long, pyramidal, secondary branches 0.6—1.8 cm
long, the rachis densely appressed furfuraceous lepidote, glabrescent, ter-
minating in 5—14-flowered corymbs; inflorescence bracts pay caducous,
membranaceous, lanceolate, 5.7—8.1(—20.2) mm long, 2.4—3.9 mm wide,
apically acute, sessile, midrib slightly raised above and a secondary
veins obscure to not visable above and below, scattered furfuraceous lepi-
dote to mostly glabrous below, densely appressed furfuraceous lepidote below,
the margins entire, ciliolate; floral bracts like the inflorescence ones but reduced
in size, 0.9-2.4 mm long, 0.2—0.8 mm wide, apically subulate; pedicels
cylindrical, 4.1-6.9 mm long, scattered, appressed furfuraceous lepidote,
usually persistent with age, persistent. F/owers 5-merous, reddish-violet; calyx
chartaceous, suborbicular, 2—2.2 mm long, the tube 1—1.1 mm long, the

—

Sipa 1834)

Fic. 5. Hymenandra squamata (Lunde IL) Pipoly & Ricketson. A. Reproductive shoot. B. Detail
ote scale from above and from the side. D. Detail of inflorescence

of abaxial leaf surface. C. Lepic

E. Detail of flower. F. Detail of stamens, showing abaxial (left) and adaxial (right) surfaces.
G. ovules. H. Fruit. A-G drawn from MO isotype, W. Burger & R. Stolze 5853. G drawn
from M. Graywm & G. Herrera 7857.

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 727

lobes almost free 1-1.2 mm long, 0.9-1.1 mm wide, asymmetric, apically
rounded, subapically notched, prominently black punctate medially, sparsely
furfuraceous lepidote, glabrescent, the margin irregular, minutely erose-fimbriate,
hyaline, densely glandular-cilioate; corolla membranaceous, 4.2—4.4 mm
long, the tube 1-1.2 mm long, the lobes connate basally, lanceolate 3—3.2
mm long, 1.2—1.3 mm wide near the base, apically acute, reflexed in an-
thesis, prominently black punctate, the margin erose, hyaline, glabrous; stamens
2.6—3 mm long; the filaments connate into a chartaceous elobate tube, the
tube 1—-1.2 mm long, adnate to the corolla basally, epunctate, glabrous; anthers
connate by their sutures, at least at first, ovate, 1.6—-1.8 mm long, 0.8-0.9
mm wide at base, sessile on the staminal tube, apically apiculate, basally
cordate, longitudinally dehiscent by narrow, sublatrorse slits, the connec-
tive prominently reddish punctate; pistil obturbinate, 2.1-3.4 mm long;
ovary 0.5—0.7 mm long, glabrous; style (1.6—)2.5—2.7 mm long, erect, in-
conspicuously black punctate, glabrous; stigma punctiform; placenta ellipsoid,
0.1—0.3 mm long, 0.1—0.15 mm in diam., apically ee ovules 7-10,
pluriseriate. Fruit depressed-globose, 4.5—6 mm long, 5.5—6.8 mm in diam,
conspicuously black punctate, the style base persistent.

Distribution.—Provinces of Heredia, Lim6n and San José, Costa Rica, at
90—1,400 m elevation.

Ecology and conservation status —Hymenandra squamata is known only from
the swamp forests of the Atlantic drainage in Costa Rica. It occurs just above
the high water level. Because these forests yield high volumes of timber,
the species should be considered under threat.

Etymology.—The epithet comes from the Latin “sgwamatus” which means
“provided with scales,” referring to the scales of the shoots and leaf blades.

Specimens examined. COSTA RICA. Heredia: Between Rio Peje and Rio Sardinalito,
Atlantic slope of Volcan Barva, 10° 17! N, 84° 04.5' W, 800—1,000 m, 7 Apr 1986 (fr), M.
Grayum & R, Chazdén 6853 (MO); Finca La Selva, the OTS Field Station on the Rfo Puerto
Viejo just E of its junction with the Rio Sarapaquf, along E River Road near Holdridge
Trail, ca. 100 m, 15 May 1978 (A), M. Graywm 1303 (DUKE); Finca la Selva, Puerto Viejo
de Sarapiqui, ne onbade E] Taconazo, Between Quebrada El Sura and W boundary
Trail, without elev., 15 Aug 1979 (fr), M. Graywm 2371 (DUKE); ee Nacional Braulio
Carrillo, 10° 15.5! 'N, 84° 05' W, 1,200-1,400 m, 13 Nov 1986 (fr), M. Grayum & G.
Herrera 7857 (MO), Finca la Selva, Puerto Viejo de Sarapaquf along Quebrada El] Santo,
Hartshorn Trail, ca. 100 m, 17 May 1980 (fl), B. Hammel 8693 (DUKE), 22 May 1980 (A),
B. Hammel 8762 (DUKE); La Selva, on Rio Puerto ee just E of its junction with Rio
Sarapaqui, Central Trail, 1,500 m S$ of edge of Trail, 100 m, 7 Jul 1981 (fr), B. Hammel
10962 (DUKE); La Selva, juse E of juction of : 1erto Viejo and Rio Sarapiquf, ca. 100
m, 20 May | = 2 (A), B. Hammel 12372 (DUKE, 7 Jun 1982 (fl), B. Hammel & J. Trainer
12757 (DUKE, LL); Finca La Selva, El Swamp 2 00 m, 16 Jun 1984 (fl), B. Jacobs 2377
(DUKE); La Selva, Sendero El Bee. ca. 2,800 m a E-W, ca. LOO m, 18 May 1985 (fl), B.
ae 3113 (DUKE); 9 km SE of San Ramin, > 16' N, 84° 05' W, 1,000 m, Sep 1985

, B. Loiselle 144 (MO); La Selva, near ee a. along line B, 90 m, 29 May 1971

~—

©C

728 Sipa 18(34)

(fl), P. Opler 188 (F), along line A, 4 Aug 1971 (ster.), P. Opler 332 (F); Finca la Selva, Puerto
Viejo, 1,600 Line Trail, without elev., 6 Jul 1979 (fr), 7. pk 790 (DUKE); Finca La Selva,
Rio Puerto Viejo just E of its junction with the Rio Sarapaqui, S Research Trail, 19 May
1984 (fl), R. Wilbur Gh, aa 34720 (DUKE), along S bounday of new property from the
SE corner to the Sarapiqui Trail at about 4,000 m on grid system, ca. 100 m, 19 May 1984
(A), R. Wilbur & B. Jacobs 34747 (DUKE); La Selva, Jaguar Trail from Quebrada El Sura to
the 3,000 m line, ca. 100 m, 14 Jul 1986 (fl, fr), R. Wilbur 39948 (DUKE); La Selva, S
Experimental Trail at the 700 m point on line; 100 m, 25 Jul 1986 (fr), R. Wilbur 40442
(DUKE); La Selva, slopes just ae out of the mp at about 200 m on the El Swampo
Trail, ca. 100 m, 29 May 1995 (A, fr), R. W “bur “GA 246 (DUKE). ae. Siquirres, Las
Brisas de Pacuarito, SOO m, ow ia 1985 (fl), L. Gomez et al. 23383 (BRIT, NY); Ridge :
of oe Pacuare, ca. 15 km S of Pacuarito, in vicinity of community of Las Brisas, 10° 00
N, 83° 30! W, 200-500 m, Jun 1988 (fl), B. Hammel & M. Chavarrfa 17073 (MO). San
ie Parque Nacional Braullio Carrillo, Carrillo station, 300-600 m, 31 May—5 Jun 1980
(fl), C. Todzia 1302 (TEX)
Hymenandra squamata is very closely related to H. sordida, but easily separated
from it by the smaller, minutely erose-fimbriate calyx lobes, the pyramidal
panicle and the dense furfuraceous lepidote tomentum, appearing velutinous.

5. Hymenandra wilburiana (Lundell) Pipoly & Ricketson, comb. nov. (Figs.

, 0). Auriculardisia wilburiana Lundell, Phytologia 63:76. 1987. Ardisia wilburiana

(Lundell) Lundell, Phytologia 63:463. 1987. Typr. COSTA RICA. Herepta: Finca

La Selva, the OTS Field Station on the Rfo Puerto Viejo juse E of junction with the

Rio Sarapiqui, Paso Perdido Trail, ca. 3,200 m line (EW), ca. 100 m, 16 Jul 1984

(A), B. Jacobs 2917 (HoLotyPE: LL; isorypes: DUKE, MO)

peat savapiquiensts Lundell, Phytol i ecmelee 1987, SYN. NOV. Ardisia Sarapiquiensis

rdell) Lundell, Phytologia 63:463. 1987. Type. COSTA RICA. Herepta: Finca La

the OTS Field Station on the - Puerto Viejo just E of its junction with the

Rio Sarapiqui, in forest on ridge at SW corner of new property, ca. 100 m, 20 Jul
1982 (A), B. Hammel & J. Trainer 13262 (HoLoryee: LL; tsorype: DUKE).

Tree 1—7 m tall, trunk to 3 cm in diam. Vegetative shoots terete, 4-6 mm
in diam., minutely scattered furfuraceous lepidote below, densely furfura-
ceous apically at least when young; reproductive shoots similar to vegetative
ones but 1.5—3.5 mm in diam. Vegetative shoot leaf blades coriaceous, elliptic
to oblong, 5.5—22.2 cm long, 1.3—7.8 cm wide, apically acuminate, the
acumen 2—3 cm long, basally acute, midrib slightly raised above, promi-
nently raised below, the secondary veins 11-19 pairs, slightly depressed or
not visible above, prominently raised below, minutely scattered furfuraceous
lepidote along the midrib below and on the blade below, glabrescent with
age, the margin entire, flat; petioles marginate, 0.5—2.2 cm long, glabrous
above, scattered and minutely furfuraceous es below; prt shoot
leaf blades as in the vegetative except 7.2-17.3 cm long, 2.1—4.7 cm wide,
secondary veins 9—15 pairs; petioles as in vegetative ones - 0.3-1.3 cm
long. Inflorescence terminal, pendent, pinnately or bipinnately paniculate, 7—
16.5 cm long, pyramidal, secondary branches 4—6.5 cm long, the rachis

PipOLy AND RICKETSON, Hymenandra and its boreotropical implications 729

. 6. Hy menandra wilburiana (Lundell) Pipoly & Ricketson. A. Reproductive shoot. B.
il of abaxial leaf surface. C. Detail of inflorescence. D. Detail of flower. E. Detail of
stamens, sae adaxial (left) surfaces. F. Detail of stamens, showing abaxial (right) sur-
faces. G. Detail of flower, showing seapls and pistil. H. ile I. Fruit. A-B drawn from
holotype, B. Jacobs 2917. C—H drawn from holotype of Auriculardisia sarapiquiensis Lundell,
B. Hammel & J. Trainer 13262. 1 drawn from R. Robles Led.

730 Sipa 18(3)

minutely scattered furfuraceous lepidote, glabrescent, terminating in 6—
| 2-flowered corymbs; inflorescence and bracts unknown (presumably
early caducous); pedicels cylindrical, 6.8—13.2 mm long, scattered furfura-

ceous lepidote, persistent. F/owers S-merous, white; calyx lobes essentially

free, chartaceous, suborbicular, 1.4—1.8 mm long, 1.6-2 mm wide, asym-
metric, notched below the apex, apically obtuse, prominently black punc-
tate medially, sparsely furfuraceous lepidote, the margin irregular, minutely
erose, hyaline, densely glandular-cilioate; corolla lobes basally free, mem-
branaceous, lanceolate, 5.5—6.5 mm long, 1.4—1.6 mm wide near the base,
apically acute, reflexed in anthesis, prominently pellucid punctate with-
out, sparsely so medially within, the margin erose, hyaline, glabrous; sta-
mens 6—6.5 mm long; filaments connate into a chartaceous clobate tube,
the tube 1.4—1.9 mm long, adnate to the corolla basally, epunctate, gla-
brous; anthers connate by their sutures, at least at first, lanceolate, 5.5—5.8

mm long, 1.9—2.2 mm wide at base, sessile on the staminal tube, apically

apiculate, basally cordate, longitudinally dehiscent by narrow, sublatrose

slits, the connective conspicuously brown or pellucid punctate; pistil obturbinate,

5.7-6.5 mm long; ovary 1.5-1.8 mm long, glabrous; style 4.2-4.7 mm

long, erect, inconspicuously pellucid punctate, glabrous; stigma punctiform;

placenta ellipsoid, 0.4—0.5 mm long, 0.2—0.3 mm in diam., apically api-

culate; ovules 16-20, pluriseriate. Frit subglobose, 7-8 mm long, 6—6.5

mm in diam., at times with noticeable longitudinal costae, densely and promi-

nently punctate, the style base persistent.

Distribution. —R fo San Juan, Nicaragua to Heredia and Limon, Costa Rica,
from O—700 m elevation.

Ecology and conservation status. —Hymenandra wilburiana 1s a ridgetop spe-
cies in the lowland to premontane wet forest transition areas. It has a rather
restricted distribution, but not enough is know of its population dynamics
to determine its conservation status.

Etymology.—The species is named for Dr. Robert L. Wilbur, professor of
botany at Duke University, expert in Mesoamerican Ericaceae and long-time
student of the flora of the La Selva OTS station in Costa Rica.

Specimens examined. NICARAGUA. Rio San Juan: Near Cafio Chontaeno, 20 km
NE of El oa 200 m, 18-21 Apr 1978 (fr), D. Nei/l & P. Vincelli 3506 (MO, NY).
COSTA RICA. Heredia: Sarapiqui Canton, trail between Canta Rana Camp and Rio Peje,
enn 100 m, 14 Jan 1983 (fr), 1. Chacin 80 (MO); Finca la Selva, on Rio Puerto Viejo
just E of its junction with Rio ae aatit Fila Rio Peje, along the line at 3,600—2,200 m,
100 m, 21-22 Apr 1983 (fr), | Chacén 717 (DUKE); N of Puerto Viejo, 10 km down road,
then 7—8 km W, without elev., 2 Feb 1983 (fr), N. Garwood et al, 936 (MO); ie El Bejuco,

S base of Cerros Sardinal, Chilamate de ps 10° 27'N, 84° 04' W, 70-100 m, 2 Jun
1985 (fr), M. Grayum & B. Jacobs 5352 (BRIT, LL, MO); S seen Canton, Pra Nacional
Braulio Carrillo, Estaci6n Magsasay, 10° oi O03" N, 84° 03! OO" W, 2 , B. Hammel et

al, 17829 (BRIT, F, MO); Finca La Selva, Sendero E] Peje, 3,200 m— ee mon line along

PipOLy AND RICKETSON, Hymenandra and its boreotropical implications 731

ridge, ca. 100 m, 28 May 1985 (fr), B. Jacobs 3189 (DUKE); eae Canton, Riau: as,
Estacién E] Plastico, 15 km de Horquetas, Sendero el Tigre, 10° 3"N, 84° 02'47" W,
700 m, 11 Jan 1994 (fr), J. Sdénchez et al. 308 (F, K); Sarapiqui ee Zona Protectora La
Selva, 6 km by road from Rio Peje crossing, 5 km SSE of Magsasay, E from oe to
ridge above Rio Peje, then S, 10° 21' N, 84° 03-04' W, 300-500 m, 14 Jan 1983 (fr), G
Schatz & M. oe 599 (DUKE); 500 m, 19 Jan 1983 (fr), G. Schatz & M. Conan 653
(DUKE, LL). Limén: SW-most ridge of Cerro Coronel, NW-facing slope, just S of the Rio
Colorado, 10° ae [ 83° 39' 30" W, 10-80 m, 17-18 Sep 1986 (fl), G. Davidse & G.
Herrera 31379 (MO), Parque Nacional Tortuguero, Estaci6n Agua Fria, ca. 7 km al SE Cerro
Azules, 10° 27! N, 83° 34' W, 70 m, 21 Jan 1988 (fr), R. Robles 1571 (MO, US); Parque

Nacional pianaction Estaci6n Agua Fria, ca. 12 cm al NE, on other side of Rfo Pueblo
uevo, over the Lomas de Sierpe, 10° 26' N, 83° i W, 80-100 m, 2 Mar 1988 (fr), R.
Robles 1666 (MO); Cerro Coronel, E of Rio . 10° 40! — 83° 40' W, 10-100 m, 24

Jan 1986 (fr), W i 23977 (MO); Cerro Coronel, E of Laguna Danto, 10° 41' N, 83°

38' W, 20-170 m, 15-20 Sep 1986 (fl), W. Stevens & O. Montiel 24432 (MO); Cerro Coronel,

along ridge from E of Laguna Danto SW toward hills E of Rio owe 10° 40-41'N, 83
38-39' W, 30-170 m, 17 Mar 1987 (fr), W. Stevens et al, 24972 (MO).

Hymenandra wilburiana is most closely related to H. pittert but may be
distinguished by the much shorter perianth and androecial parts, smaller
leaves and much thinner stems.

Populations corresponding to the type of Avriculardisia sarapiquiensis are
notable only for their slightly smaller inflorescence and slightly smaller leaves.
The types of Hymenandra wilburiana and Auviculardisia savrapiquiensis were
both collected at the Finca La Selva OTS Station.

6. Hymenandra pittieri (Mez) Pipoly & Ricketson, comb. nov. (Figs. 1C,
7). Ardisia pittieri Mez, Bull. va oe Ser. 2, 3:236. 1903. Auriculardisia pittieri
(Mez) Lundell, Phytologia 49:345. 1981. Type. COSTA RICA. Cartaco: Las Vachs,
Tucurrique, ease 700 m, May | oe ee a. Tonduz 13369 (LEcroTyPE here desig-
nated: US (US # 1339917, LL Neg. # 1971-83); isoLecrorypEs: G 4-sheets, P 4-sheets,
US (US # 1339916, LL Neg. # 197 1-83)).

Ardisia cuttert Standl., J. Wash. Acad. Sci. 17:52. 1927, syN. Nov. Auriculardisia cutteri

(Standl.) Lundell, Wrightia 7:267. 1984. Type. COSTA RICA. Ses vicinity of
Pejivalle, ca. 900 m, 7 Feb 1926 (fr), P. Standley & J. Valerio 47122 (HoLotTyPE: US,
(US Neg. # 2368, LL Neg. # 1971-31).

Ardisia coclensis Lundell, Ann. Missour Bot. Gard. 28:453. 1941, syn Nov. Auriculardisia
coclensis (Lundell) andell, Phytologia 49:343. 1981. Typr. PANAMA. Coc Le: vicin-
ity of El Valle de Antén, ca. 600 m, without date (A), P A//en 2056 (HoLotyPE: MICH,
(LL Neg. 4 1971-4)). [Unicate].

Tree (0.9—)2—5(—8) m tall, 2-8 cm in diam. Vegetative shoots terete, 7-24
mm in diam., minutely scattered furfuraceous lepidote, glabrescent; repro-
ductive shoots as in the vegetative but 2-9 mm in diam. Vegetative shoot leaf
blades coriaceous, obovate to oblanceolate, 33.7—72.2 cm long, 5.3—16.5
cm wide, apically acute to rounded, gradually tapering to a cuneate base,
decurrent to near the petiole base, midrib slightly raised above, prominently
raised below, secondary veins 24—41 pairs, prominently raised above and

Sipa 18)

~~
wD
No

Fic. 7. Hymenandra pittieri (Mez) Pipoly & Ricketson. A. Reproductive shoot. B. Detail of
abaxial leaf surface. C. Detail of inflorescence. D. Detail of flower. E. Detail of stamen, showing
abaxial (left) and adaxial (right) surface. F Ovules. G. Fruit. A—B drawn from G. de Nevers
7203. C, F—G drawn from P lectotype, A. Tonduz 13369. D-E drawn from J. Morales & Y.
Salazar 1195.

Pirpoty AND RICKETSON, Hymenandra and its boreotropical implications ee,

below, prominately punctate above and below, glabrous above, minutely
scattered furfuraceous lepidote below, glabrescent, the margin regular, en-
tire, flat; petioles marginate 1.84.7 cm long, glabrous above, minutely scattered
furfraceous-lepidote below; reproductive shoot leaf blades similar to the veg-
etative but 3.9-34.3 cm long, 1.2-11.8 cm wide, secondary veins 20—36
pairs, basally tapering to a rounded base appearing auriculate; petioles similar
to the vegetative ones but 0.4—1.1 cm long. Inflorescence subterminal, pin-
nately or bipinnately paniculate, (7—)12—30(—36) cm long, pyramidal, sec-
ondary branches 3.5—12.5 cm long, the rachis densely and minutely furfu-
raceous tomentose, terminating in 8—13-flowered corymbs; inflorescence
bracts early caducous, membranaceous, oblong, 7—13.5 mm long, 3.4—4.6
mm wide, apically acute to rounded, basally sessile, midrib slightly raised
above and below, secondary veins not visible, prominently punctate, densely
and minutely furfuraceous tomentose, the margin entire, flat; floral bracts
1.7—2.8 mm long, 1.1—2.1 mm wide, otherwise as in the inflorescences bracts;
pedicels cylindrical, 7.5-14 mm long, densely and minutely furfuraceous
tomentose. Flowers S-merous, pink; calyx lobes almost free, chartaceous,
suborbicular, 2.2—-2.8 mm long, 2—2.4 mm wide, asymmetrical, subapi-
cally notched, apically obtuse to broadly rounded, prominently punctate,
densely and minutely furfuraceous tomentose, the margin, irregular, mi-
nutely erose, hyaline, sparsely glandular-ciliolate; corolla lobes basally free,
chartaceous, ovate to lanceolate, 9-10.5 mm long, 44.7 mm wide, apically
acute, reflexed in anthesis, prominently pellucid punctate on both surfaces,
the margin hyaline, entire, glabrous; stamens 8.7—9.5 mm long; filaments
connate into a coriaceous elobate tube, the tube 1.2—1.5 mm long, free from
the corolla basally, epunctate, glabrous; anthers connate by their sutures, at
least at first, lanceolate, 7.5—8 mm long, 1.4—1.7 mm wide at base, sessile
on the staminal tube, apically acute, at times with a minute emarginate
tip, basally cordate, longitudinally dehiscent by narrow, sublatrose slits, the
connective inconspicuously pellucid to black punctate; pistil 6.8—8.4 mm
long; ovary 1.8—2.2 mm long, glabrous; style 5—6.2 mm long, erect, in-
conspicuously punctate, glabrous; stigma punctiform; placenta ellipsoid,
0.9-1.1 mm long, 0.8-0.9 mm in diam., apically apiculate; ovules 14—16,
pluriseriate. Fruit globose, 0.7—1.6 cm in diam., densely and prominently

lack punctate, the style base persistent, the exocarp thick, juicy, bright red.

Distribution —Limo6n, Cartago, San José and Puntarenas, Costa Rica and
Coclé, Colon and San Blas, Panama, from 50—1,300 m elevation.

Ecology and conservation status —Hymenandra pittieri is an understory tree
along ridgetops in lowland and premontane wet forest. Throughout its range,
this species is locally common and is not threatened at this time.

Etymology.—This species is named for Dr. Henri Pittier, early explorer in

734 Sipa 18(@)

Central America but also known for his work in South America and par-
ticularly, for establishing the Herbario Nacional de Venezuela.

ewe examined. COSTA RICA. Cartago: Along Camino Raiz de Hule, SE of Platanillo
(Tsipirf), 1,200-1,400 m, 1 Jul 1976 (A), 7. Croat 36703 (LL, MO); 24km NE of Turrialba
on hwy. to Limon, then E at Tres Equis on ha road 1.5 km, 09° 58' N, 83° 34' W, 450-
525 m, 10 May 1983 (fl), R. Liesner et al. 15348 (MO); Pavenes. Turrialba, 650 m, 18 May
1972 (fA), L. Poveda 106 (MO). Limon: Near the Rio Catarata (Rio Sand ce in the hills
between Bri me on the Rfo Sixaola and the Caribbean ee plain, 09° 37! N, 82° 49! W.

’

50-100 m, 28-29 Nov 1975 (fr), R. Baker & W. B 26 (F). Puntarenas: Cantén de
Osa, Ret ae Nacional Golfo Dulce, Peninsula de oF fey Chal, entrance to Chocuaco,
O8° 43! 00" N, 83° 34' 50" W, 200—350 2 Nov 1994 (fr), R. Aguilar et al. 36063 (MO);

ie between Rio Esquinas & Palmar ce de Osa, sea level, 30 Jan 1951 (fr), P. A/len
5828 (F), 75 m, 2 Feb 1951 (fr), P Aflen 5828A [sic] (F), 60 m, 18 Feb 1953 (fl), P A//en
6726 (F); Canton de Coto Brus, P.I. La Amistad, Cordillera de Talamanca, Las Cruces, 08°
47' 20" N, 82° 58' 30" W, 1,200-1,300 m, 30 Jun 1995 (fr), L. Angulo 394 (MO); For-
ested slopes E of Las Cruces and 5—6 - S of San Vito on and around the property of Mr.
Robert Wilson, 8° 47' N, 82° 58' W, 1,100-1,200 m, 15-16 Jan 1967 (fr), W. Burger &
G. Matta U. 4406 (F); Oeioe forest ca. : ) km SE of Palmar Norte along the Interamerican
Hwy., to Panama, 08° 54' N, 83° 02' W, 20 m, 26 Jan 1967 (fl), W. se &G. Matta U.
4644 (F, G); Canton de Osa, Rincén, Fila Casa Loma, 08° 43' N, 83° 32' W, 50-400 m, 22
Jul 1990 (fr), A. Chacin 934 (MO); Quebradas Pastora, Arepa y Scie Fila Esquinas-
Osa, without elev., Nov 1983 (fr), 1. C babe On i G. Herrera 1758 (MO); Along Rio Jaba S of
San Vito de Coto Bry, 08° 47' N, 82° 58' W, 1,150 m, | Jul 1984 (fr), M. Grayum et al.
3369 be Refugio Nacional Golfito, both slopes of . ila Gamba « . along crest of same,
to ca. 0.7 km N of Golfito- Villa Biieedo road, O8° 40.5' N, 83° , 160-260 m, 11
Dec 1988 aes G&G G. Herrera 9200 as egene ee to e hacarita, along
Interameric ., ca. 2 km N of Chacarita, 08° 48' N, 83° 18' W, 50 m, 25 May 1986
(fr), B. ey et a 15186 (MO); Alto La Palmera, Carretera nee e 950 m, 28
Feb 1966 (fr), A. cle (DUKE, F); Cantén - oar Rancho Quemado, sector E, Sierpe,
08° 40' 20" N, 83° 35' 15" W, 500 m, 6 Nov 1991 (fr), J. Marin et al. 257 (MO); Finca las
Cruces, along ee to stream below ne field station, without elev., 10 May 1973 (fl), B.
McAlpin 2250 (DUKE); Cantén de Osa, Camino a la toma de agua, Rancho Quemado Rincén,
08° 42'N, 83° 34' W, 200 m, 9 Dec 1990 (fr), F Quesada 320 (MO); Wilson’s any 6km
S of San Vito de Java, 4,000 ft [1,219 m], 16 Aug 1967 (fr), P. Raven 21829 (F 2-sheets,
MO); Golfito, Parque Nacional Esquinas, margins of Quebrada Gamba, 08° 40! 30" N,
83° 12' 20" W, 70 m, 29 Sep 1995 (fr), J. Séuchez 565 (CR, F). San José: Canton de Pérez
Zeledén, Tinamaste, Swiss farm, 09° 17' 40" N, 83° 46' 00" W, 1,000 m, 12 Jun 1997 (fr),
D. Argiiello M. et al. 11 (CR, F, MO, NY); Valley of the Rio Hondura, below La Palma, NE
Be San Jeronimo, 10° 3' N, 83° 58' W, 1,000 m, 15 May 1968 (A), W Burger G R. Stolze
38 (FE, MO); El General Valley, a of San Isidro El General, Alto San Juan, road to
sense 900 m, 28 Feb 1966 (A, fr), A. Molina R. et al, 18090 (F 2-sheets, NY, US);
Z.P. La Cangreja, Santa Rosa de Puriscal, faldas de la Fila La Cangreja, 09° 42' 28" N, 84°
23' 38" W, 500 m, 8 Mar 1993 (A), J. Morales & Y. Salazar 1195 (MO); Canton de Acosta,
Fila Bustamante, by Fila San Jerénimo, headwaters of Quebrada Colorado, 09° 43' 20 N,
84° 16' 45 W, 1,040 m, 29 May 1994 (fr), J. Morales & V. Urena 2852 (INB-2 sheets, MO
2-sheets); Cant6én + Amidrib, Fila Bustamante, Zoncuano, a r de Fila Zoncuano, ca.
del Rio Parritilla, 09° 42' 02" N, 84° 13' 37" W, 700 —900 m, A Jun 1995 (fr), J. Morales
4354 (MO); Zapaton de Puriscal, 400 m, 1 Aug 1986 (fr), N. Ge 0 Jiménez 1208
(MO). PANAMA. Coclé: Hills N of El Valle de Antén, 1,000 m, 23 Jun 1940 (fr), P.

pace,

PipOLY AND RICKETSON, Hymenandra and its boreotropical implications 13)

Allen 2176 (F); Loma del Tigre, region N of El Valle de Antén, 1,000 m, 16 Nov 1946 (fl),
P. Allen 3806 (G, MO, NY); 7 km from Llano Grande on road to Coclesito near continen-
tal divide, 1,200 ft [366 ml], Jul 1979 (fr), 7; Antonio 1364 (MO); On the Atlantic side, ca.
5 hr. walk from sawmill at E] oie along slopes above Norte Rio Blanco near small vil-
lage of Cafio Sucio, 400-500 ft [122-152 ml], 2 Feb 1980 (fr), T. Antonio 3627 (LL, MO),
3632 (MO); Area of El Valle, 2 km : of La Mesa, N slope of Cerro Gaital, 08° 38' N, 80°
07' W, 800 m, 16 Nov 1983 (fr), H. Churchill 3867 (LL, MO, NY); 3876 (MO); Near
continental divide along lumber road ca. 1.5 mi N of El Cope, ca. 900 m, 19 Jan 1978 (fr),
T. Croat 44581 (BRIT, MO); Hills above E] Valle de Antén, without elev., 13 Aug 1972
(fr), W. D’Arcy & J. D’Arcy 6749 (MO); Road from Penonome to Coclesito, 9 km N of
Llano Grande, on tributary on Rio Caseaja, without elev., 11 Oct 1978 (fr), W. D'Arcy G B.
Hammel 12292 (LL, MO); Area between Cafio Blanco dg Norte, Cafio Sucio and Chorro
del Rio Tife, 08° 43' 06" N, 80° 36' 30"—38' 00" W, 200-400 m, 3 Feb 1983 (fr), G.
Davidse & C. Hamilton 23484 (MO): Hills N of El Valle de Antén, ca. 800 m, 17 Jan 1973
(fl, fr), R. Dressler 4254 (MO); Near Aserradero El Copé, N of El Copé, E of sawmill, 700—
800 m, 2 Nov 1980 (A, fr), R. Dressler 5962 (FLAS 2-sheets, MO); Slopes of Cerro Pilén
near El Valle, 700—900 m, 10 Jun 1967 (fr), J. Duke 12197 (MO); Summit of Cerro Pilén,
above El Valle de Antén, 2,700 ft ee ml}, 28 Mar 1969 (fr), J. Dwyer et al. 4476 (LL,
MO); Cerro Pilén, bottom of slope, ca. 2,500 ft [762 m], 19 Jan 1968 (fr), J. Dwyer 8351
(MO), 13 Jul 1968 (fr), J. Dwyer & “7 Lallathin om (EF, MO), 14 Jul 1968 (ster.), SO90A
(MO); Adjacent to chicken farm, La a, above El Valle, without elev., 3 Jan 1974 (fr), J.
Dwyer areee 9 (MO); 7 km N of el Copa “ Veraguas, near Bete sawmill, Alto Calvario,
900-1,300 m, 11 Jan 1977 (fl), J. Folsom 1242 (BRIT, LL, MO); New Works at Rivera
sawmill, Alto Calvario, (00-800 m, 12 May 1977 (fr), J. Folsom 3163 (MO); Area around
Rivera sawmill, 7 km N of El Cope, Atlantic slope, N side of summit, Alto Calvario, 700—
850 m, 3 Oct 1977 (fr), J. Folsom et al. 5741 (MO); Vicinity of La Mesa, N of El Valle,
1,000 m, 23 Dec 1972 (fr), A. ee 6854 (LL); Hills above El Valle, ca. 1,000 m, 24 Dec
1972 (fl), A. Gentry 6872 (LL, MO); Near continental divide along lumber road, 8.4 km

above E] Cope, 1 km beyond ee 900 m, 19 Jan 1978 (fr), B. Hammel 958 (LL
7 km N of Llano Grande on road to Coclesito, 1,700 ft {518 m}, 8 Mar 1978 (fr), B. ee,
1928 (MO); La Mesa region N of Cerro Gaital vicinity of El Valle, 2,400 ft [732 ml}, 2 Jul
1978 (fr), B. Hammel 3898 (MO); Continental divide N of Penonome on road to Coclesito,
1,600 ft [488 m}, 25-26 Jul 1978 (fr), B. pig 4054 (MO); S of Cascajal along Conti-
nental Divide, 8° 45' N, 80° 25' W, 800-900 m, 7 Nov 1981 (fr), 8. Knapp 1965 em
Cerro Pilén, 2,700 ft [823 ml], Jul 1968 (fr), B. Lellthin 35A (MO); Along road ca. 8
N of El ie de Anton, without elev., 3 Aug 1970 (fr), J. Lateyn G H. Kennedy 1704 (DUKE,
GH, LL, MO); Vicinity of la Mesa, beyond El V ae N slopes of Cerro Gaital, 8° 37'N
80° 07! ee 850 m, 14 Jul 1987 (fr), G. McPherson 11270 (MO); Ca. 3 km NE of an Ville.
along farmers road, without elev., 2 Nov 1974 (Al, fr), 8. Mori & J. Kallunki 2978 (LL, MO);
Continental Divide above El Copé, 8° 38' N, 80° 39' W, 650-750 m, 27 Nov 1985 (fl), G.
de Nevers et al. 6398 (LL, MO); Foot of Cerro Pil6n, above El Valle de Antén, 2,000 ft [610
ml}, 27 Mar 1969 (fr), D. Porter et al. 4422 (MO), 28 Mar 1969 (fr), D. Porter et al. 4597
(LL, MO); La Mesa, 2 km NW of Cerro Pil6n, 800 m, 22 Jul 1976 (fr), G. Su/livan 553 3
(MO); Between Rio Blanco and Cafia Susio 1 hr. hike to the W, Rio Blanco is ca. 5 hr. h
N down from Continental Divide above El Copé and El P so sawmill, 8° 38'N, 80° 36)
W, 350-400 ft [107-122 ml], 13 Dec 1980 (fr), K. ae et al, 2453 (LL, MO); Ca. 1 km
E of Quebrada Amarillo in La Mesa, 2.5 km N of El Valle, 8° 37! N, 80° 07' W, 2,600 ft
{792 m}, 14 Feb edi (fr), K. Sytsma & W. D’Arcy 3565 (LL, MO). Colén: Santa Rita lum-
ber road, ca. 15 km E of Colén, without elev., 5 Oct 1969 (A, fr), R. Dressler & W. Lewrs
3725 (LL, MO); ae Rita Ridge, logging area 19 km in from Transisthmian Hwy., with-

736 Sipa 18(3)

out elev., 28 Jan 1968 (fr), J. Dwyer 8551 (F, MO); Rio Escandaloso near the abandoned
Manganese mine, 500 ft [152 m]}, 26 Apr 1978 (fr), B. Hammel 2662 (LL, MO); S approach
to Cerro Bruja from Rio Escandaloso, ridge top, without elev., 20 May 1978 (fr), B. Hammel
3220 (MO); On Santa ee Ridge Trail, beyond end of Santa Rita Ridge Road (Panama-

nian Hwy. R2OD), 17-35 km from Boyd-Roosevelt Hwy, 400-800 m, 21 May 1975 (fr),
S. Mori & M. Crosby 6313 (LL, MO); Santa Rita Ridge, me 13.8, 9° 20'N, 79° 45! 7 350
m, 24 Feb 1986 (fr), G. de Nevers 7203 (LL, MO). San Blas: Rio Cangandi, pueblo Cangand1,
camino de caballos al S del pueblo, 9° 27' N, 79° 07' O, 40 m, 17 May 1987 (fr), H. Herrera
G P. Perez 98 (MO); campamento Nusagandi, en la Carretera El Llano-Carti, a 19.1 km
la Carretera Panamericana, Cuadrante Nusagandi y Sendero Nusagandi, 9° 12' N, 78° 16
W, 350 m, 19 Mar 1993 (fr), R. Paredes 944 (BRIT, EF MO, PMA, STRI).

Hymenandra pittieri is most closely related to H. wilburiana, but is a much
more robust plant, with larger sepals, stems and androecial parts.

Populations corresponding to the type of Ardisia cutteri are notable ac-
cording to Standley for their large leaves and fruits. The subsessile leaf bases
appearing auriculate have also been used to recognize it. However, we have
now found that the subsessile leaves are restricted to the flowering shoots
of Hymenandra pittieri, so the confusion is the result of an incomplete speci-
men rather than due to biological difference. Lundell (1971) stated that
Ardisia coclensis may have affinity to A. pittier7, but he only knew it from the
brief description by Standley (1938).

is gaa callejasii (Pipoly) Pipoly & Ricketson, comb. nov. (Figs.
8). Ardisia callejasii ae, Novon 2:389. 1992. Type. COLOMBIA. ANTIOQUIA:
wane de Mutata, N of Hacienda El Darién, right margin of Rfo Chontadural,
see elev., 28 Jul 1978 (A. a R. Fonnegra G. & FE. Renteria A. 953 (HOLOTYPE:

UA; tsorypes: COL, MEDEL, MO).

Tree to 7 m tall. Vegetative shoots terete, 7-10 mm in diam., densely and
minutely furfuraceous lepidote; reproductive shoots unknown. Vegetative shoot
leaf blades coriaceous, oblanceolate, (25—)31—45.5 cm long, (S—)7—12.2 cm
wide, apically long-attenuate, the acumen |.8—3.5 cm long, gradually ta-
pering to a cuneate base, decurrent to petiole base, midrib slightly raised
above, prominently raised below, secondary veins 16—20 pairs, slightly de-
pressed or not visible above, prominently raised below, glabrescent, the margin
regular, entire, flat; petioles marginate (0.6—)1—1.2 cm long, glabrous above,
densely and minutely furfuraceous lepidote below; reproductive shoot leaves
unknown. Inflorescence subterminal, pinnately or bipinnately paniculate, (7.5—)

12.5-24.5 cm long, pyramidal, secondary branches 3—12 cm long, the ra-
chis minutely densely furfuraceous lepidote, glabrescent, terminating in 7—
12-flowered corymbs; inflorescence bracts unknown (presumably early ca-
ducous); floral bracts early caducous, membranaceous, ovate to oblong, 2.3—2.5
mm long, |.2—1.4 mm wide, apically acute to rounded, sessile bases, mid-
rib slightly raised above and below, secondary veins not visible, prominently
punctate, scattered and minutely furfuraceous lepidote, the margins erose,

PiIpoLy AND RICKETSON, Hymenandra and its boreotropical implications

Fic. 8. Hymenandra eee ae : Pipoly & Ricketson. A. Reproductive shoot. B. De-
irfac

tail of abaxial leaf su section of pistil and calyx, is te calyx lobes,
elongate style, and eo. ce D. Longisection of pistil, ee . pla-
ns. F.

centa pare to ovules. E. Detail of flower, showing oor lobes and stam
A-F drawn from holotype, R. Fonnegra G. & E. Renteria A 953. Original deauiee

os Bipol (1992

hyaline, entire, flat; pedicels cylindrical, 0.7—1.8 cm long, densely furfura-
ceous lepidote, persistent. F/owers 5-merous, pink; calyx lobes almost free,
chartaceous, ovate, 2—2.8 mm long, 1—1.3 mm wide, symmetrical, apically
obtuse, prominently punctate medially, densely and minutely furfuraceous

738 Stipa 18(3)

lepidote, the margin irregular, erose, hyaline, sparsely glandular-cilioate;
corolla lobes basally connivent, coriaceous, ovate, 5.3—5.9 mm long, apically
acute, highly reflexed in anthesis, medially thickened, prominently pellu-
cid punctate without, somewhat so medially within, the margin hyaline,
entire, glabrous; stamens 4.9—5.4 mm long; filaments connate into a coria-
ceous elobate tube 1.2 Ht 3 mm long, epunctate, glabrous; anthers free, lanceolate,

—4,.2 mm long, 1.3—1.6 mm wide, sessile on the staminal tube, apically
attenuate, basally ee. longitudinally dehiscent by narrow, sublatrose
slits, che connective inconspicuously pellucid punctate; pistil ellipsoid; ovary
—4 mm long, erect, inconspicuously

8

1.5—1.7 mm long, glabrous; style 3.8
pellucid punctate, glabrous; stigma punctiform, placenta ellipsoid, 0.8—1
mm long, 0.3—0.4 mm in diam., apically apiculate; ovules 12-15, pluriseriate.
Fruit globose, 8-10 mm in diam., densely and prominently black punc-
tate, the style base persistent.

Distribution —Endemic to the Colombian Chocé Floristic Province, in the
Municipio de Mutata, Department of Antioquia, from 80-180 m elevation.

Ecology and conservation status.—This species is locally common on one farm,
in lowland pluvial forest, but has not been found elsewhere. Therefore it
should be considered threatened. The lowland pluvial forests of the Colombian
Chocé are some of the wettest in the tropics, receiving annual precipitation
well in excess of 8,000 mm per year.

Etymology —Vhe species was named in honor of Ricardo Callejas Posada,
head of the graduate program in biology at the Universidad de Antioquia,
Medellin, Colombia. Ricardo is the pre-eminent authority on the phylog-
eny and systematics of the Piperaceae on a worldwide basis.

Specimens se . OLOMBIA. Antioquia: 5 ee ecaue Longani, del Rio Longani,

2 kms. N de Mutatd, 7° 20' N, 76° 30' O, 80-10 19 Nov 1987 (fl), R. Callejas et al.
5683 (NY); Sere ae margin of Rio ie aa Villa Arteaga, Las Caucheras,
hill above water tap, 10 m, 2 Oct 1961 (bud), J. Cuatrecasas 26156 (US); Hacienda

El Darién, right bank : es ee , 20 Jun 1979 (fr), R. Fonnegra et al. 1228 (COL,
HUA, MO), (fl) 1249 (COL, HUA, MO), 13 Sep 1979 (fr), R. Fonnegra et al. 1343 (COL,
HUA, MO).

Hymenandra callejasii is rather isolated within the genus; its inflorescence
is rather unusual, as is the fact that no leaves have been seen on reproduc-
tive shoots. Further field study will be necessary to document the entire
life history of Hymenandva callejasit.

8. Hy aa acupsouns (Cuatrec.) Pipoly & Ricketson, comb. nov. (Figs.

1G, 9). P s Cuatrec., Revista Acad. Colomb. Ci. Exact 8:324. 1951
Pa arainin uatrec. = ipadall Wrightia 4 1:53. 1968. Type. COLOMBIA. Var
[pe Cauca}: Costa del Pacifico, Rio Cajambre, Barco, 5-80 m, 28 Abr 1944 (fr), J.
Cuatrecasas 17219 (WOLOTYPE: F; lsorypes: LL, US).

Tree 2-6 m tall. Vegetative shoots terete, 7.5—9.5 mm in diam., sparsely

PIpOLY AND RIcKETSON, Hymenandra and its boreotropical implications tog

and minutely meta OUs lepidote, glabrescent; reproductive shoots as in the
vegetative ones but 2.5—3.5 mm in diam. Vegetative shoot leaf blades coria-
ceous, ovate to bncee late: 15.4-27.2 cm long, 9.3-10 cm wide, apically
acute to long-attenuate, the acumen when present 0.5—1.5 cm long, ba-
sally acute to cuneate, midrib slightly depressed above, prominently raised
below, secondary veins 18—36 pairs, slightly depressed or not visible above,
raised below, glabrous throughout, the margin entire, flat; canaliculate, 2.1—
2.3 cm long, minutely appressed furfuraceous lepidote, glabrescent; repro-
ductive shoot leaf blades as in the vegetative ones except 15.5—17.8 cm long,

.8 cm wide, the secondary veins 18—32; petioles as in the vegetative
ones but 2.1-2.5 cm long. Inflorescence terminal, pinnately or bipinnately
paniculate, 9-16.5 cm long, obpyramidal, secondary branches 5—5.5 cm
long, rachis glabrous throughout, terminating in 6—12-flowered corymbs;
inflorescence and floral bracts unknown (presumably early caducous); pedicels
cylindrical, 0.9-1 cm long, glabrous, persistent. F/owers 5-merous; calyx
lobes almost free, chartaceous, oblate, 1-1.1 mm long, 1.4—1.6 mm wide,
symmetrical, apically obtuse, prominently black punctate medially, glabrous,
the margin irregular, erose-fimbriate along entire length, hyaline, sparsely
glandular-ciliolate; corolla, stamens and pistil unknown. Fruit globose, 8—
12 mm in diam., densely and prominently black punctate, the style base
persistent.

Distribution.—From the central Chocé Floristic Province of Colombia,
from 5—80 m elevation.

Ecology and conservation status.—This species occurs in coastal forests along
rivers, an area frequently colonized by squatters. Therefore, the species should
be considered threatened.

Etymology —The epithet is the superlative form of the Latin word, “acutus,’
referring to the apically acute to long-attenuate leaf apices.

Specimens examined. COLOMBIA. Chocé6: Rio San Juan Basin, Rio Bicordé, above
Noanamié; river margin; 04° 42' N, 76° 55' W, without elev., 6 Apr 1979 (fr), E. Forero et
al. 4713 (COL, MO).

Hymenandra acutissima is most closely related to the vicariant H. crosbyz,
but easily recognized because of its much longer sepals, terminal inflorescence
and much larger fruit.

); a ere crosbyi (Lundell) Pipoly & Ricketson, comb. nov. (Figs.

10). Ardisia crosbyi Lundell, Wrightia 6:73. 1979. Icacorea crosby; (Lundell) Lundell

Pl 49:348. 1981. Typr. PANAMA. CoLon: On trail at end of Santa Rita

Ridge Road, 18—30 km from Boyd—Roosevelt Highway, 600-800 m, 30 May 1975

(fr), S. a & M. Crosby 6423 (HoLotyPE: LL (F Neg. # 55657); isorype: MO (LL
Neg. # 1979-1)).

Ardisia calvarioana Lundell, Wrightia 6:66. 1979, syN Nov. Icacorea calvarioana (Lundell)
Lundell, Phytologia 49:347. 1981. Type. PANAMA. Coc if: Around Rivera Sawmill

740 Sipa 18)

Fic. 9. Hymenandra acutissima (Cuatrec.) Pipoly & Ricketson. A. Reproductive shoot. B.
Detail of abaxial leaf surface. C. Inflorescence. D. Fruit. A-B & D drawn from holotype, /.
Cuatrecasas 17219, C drawn from isotype, J. Cuatrecasas 17219.

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 741

near to Alto Calvario, 7 km N El Cope, New Works, trout stream, 700-850 m, 20
Oct 1977 (fr), J. Folsom & R. Page 5970 (HoLoryPe: MO; isoryre: LL).

Tree 1-3 m, 1-3 cm in diam. Vegetative shoots terete, 3-5 mm in diam.,
densely and minutely furfuraceous lepidote when young, glabrescent; re-
productive shoots as in the vegetative ones but 1.5—3 mm in diam. Vegetative
shoot leaf blades chartaceous, ovate or lanceolate to elliptic, 11.5—22 cm long,

—7.6 cm wide, apically long-attenuate, the acumen 1.2—2.4 cm long,
basally obtuse to rounded, only slightly decurrent, midrib slightly raised
above, prominently raised below, secondary veins 12-30 pairs, slightly de-
pressed above, or not visible above, prominently raised below, prominently
pellucid to black punctate above and below, glabrous above, scattered and
minutely furfuraceous lepidote below, the margin entire, flat; petioles mar-
ginate, 1.7—-5.4 cm long, glabrous above, scattered and minutely furfura-
ceous lepidote below; reproductive shoot leaf blades as in the vegeative ones
except 5.5—-15.7 cm long, 1.7—5.2 cm wide, secondary veins 10—26 pairs;
petioles like the vegetative ones except 1.6—3.9 cm long. Inflorescence sub-
terminal, pinnately or bipinnately paniculate, 2.7-8.8 cm long, obpyramidal,
secondary branches 1.3—3.5 cm long, rachis sparsely and minutely furfura-
ceous lepidote, early glabrescent, terminating in 4—6-flowered corymbs;
inflorescence bracts membranaceous, lanceolate to elliptic, 1.4—2.4 mm long,
0.4—0.7 mm wide, apically acute, pellucid to black punctate medially, scattered
and minutely furfuraceous lepidote below, the margin entire; floral bracts
like the inflorescence bracts but 0.8—2.1 mm long, 0.4—0.8 mm wide; pedicels
cylindrical, 0.8-12.2 cm long, scattered and minutely furfuraceous lepi-
dote, glabrescent, persistent. F/owers 5(—6)-merous; calyx lobes (in fruit) almost
free, chartaceous, ovate, 0.9-1.6 mm long, 0.9-1.2 mm wide, symmetri-
cal, apically acute, conspicuously pellucid to black punctate, scattered and
minutely furfuraceous lepidote, glabrescent, the margin irregular, erose, hyline,
glandular-cilioate; corolla 3.5—3.7 mm long, membranaceous, lanceolate,
the tube 0.3—0.5 mm long, the lobes (in young bud) basally connate, 3.1—
3.3 mm long, 1.2—1.4 mm wide at base, apically acute, prominently pellu-
cid punctate, somewhat so medially within, the margins erose, hyaline, glabrous;
stamens (in young bud) 2.6—2.7 mm long; filaments connate into a mem-
branaceous elobate tube, the tube 0.3—0.5 mm long, adnate to the corolla
basally, epunctate, glabrous; anthers connate by their sutures, at least at
first, lanceolate, 2.2-2.5 mm long, 0.4—0.6 mm wide at base, sessile on staminal
tube, apically apiculate, basally cordate, longitudinally dehiscent by nar-
row, sublatrose slits, the connective prominently pellucid aa pistil

—3.2 mm long; ovary 0.7—0.9 mm long, glabrous; style 2.1—2.5 mm long,
erect, inconspicuously pellucid punctate, paren: stigma ee ovules
22-24, pluriseriate. Fruit globose, 4.5—6.5 mm in diam., prominently raised
pellucid to black punctate, glabrous, the style base persistent.

742 Stipa 18(3)

Fic. LO. Hymenandra crosbyi (Lundell) Pipoly & Ricketson. A. Reproductive shoot. B. De-
cail of oo leaf surface. C. Detail of inflorescence. D. Detail of calyx and pedicel. E.
Fruit. A-B & D drawn from the holotype, $. Mori & M. Crosby 6423. C drawn from G. de
Nevers 6757. E drawn from K. Sytsmea et al, 427°

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 743

Distribution —Restricted to the Caribbean slope, in the provinces of Coclé,
Col6n and San Blas, Panama, from 100—850 m elevation.

Ecology and conservation status —H ymenandra crosbyi occurs at the natural
ecotone of premontane and cloud forest. Its conservation status cannot be
determined owing to a lack of populational data.

Etymology.—The species is named for Dr. Marshall Crosby of the Mis-
sour! Botanical Garden, who collected the type specimen.

Specimens examined. PANAMA. Coclé: Coclecito oer elevational transection from

1 mile beyond the divide to the ridge top, 08° 42' N, 80° 28' W, 300-500 m, 12 Jan 1986
(fr), G. de Nevers et al. 6757 (LL, MO). Colén: ae oe ene headwaters, along trail
from end of Santa Rita Ridge Road, ca. 11 km SW of Cerro Braja, 09° 25' N, 79° 35' W,

600-700 m, 2 May 1981 (fr), K. Sytsma et al. 42 (LL, MO); Ridge between Rio Sie
and Rio Gatun waterbeds, any Me from end of Santa Rita Ridge road, ca. 5-8 km SW
of Cerro Bruja, 09° 27' N, 79° 36' W, 700-800 m, 2 May 1981 (fr), K. Sytsma et al, 4279
(LL, MO). San Blas: El eee Road, 17.4 km from Interamerican Hwy., 09° 19'N,
78° 55' W, 350 m, 27 Sep 1984 (fr), G. de Nevers et al. 3941 (MO); Trail along Continental
Divide, 5-10 km W of El Llano-Carti Road, 350 m, 10 Jan 1985 (fr), G. de Nevers & H.
Herrera 4490 (MO), Rio Nergala, 9° 22'N, 79° 07' W, 100-300 m, 12 Jan 1985 (fl), G. de
Nevers G H. Herrera 4531 (MO)

Hymenandra crosbyi is most closely related to the vicariant H. acutissima,
but may be recognized by the shorter and obtuse sepals, the subterminal
inflorescence, shorter secondary branches, and smaller fruit.

ACKNOWLEDGMENTS

We thank the Missouri Botanical Garden and the Flora Mesoamericana
Project for funding that allowed J. Ricketson (MO) to travel to Fort Worth
and for J. Pipoly to travel to MO. We have enjoyed the continuing support
of the staff of the C.L. Lundell Herbarium (LL), housed at the University of
Texas at Austin. Without access to that critical collection, assembled by
C.L. Lundell over a period of nearly 60 years, the present study would not
have been possible. We also thank the curators of the herbaria cited for loans
of specimens. We are grateful to the staff of TEX/LL, especially Billie L.
Turner, Tom Wendt, Carol Todzia, Beryl Simpson, and José Panero, for their
cooperation and hospitality. We are also grateful to those who have been so
instrumental in assisting us in our work, including Gerrit and Jeany Davidse,
Linda Oestry, Mary Bard, and Catherine Mayo (MO), and Barney Lipscomb
and Lindsay Woodruff (BRIT). Illustrations were prepared by the junior
author except for H. callejasiz, drawn by Juan Pinz6n (COL). Reviews of the
manuscript by Tom Wendt (TEX), and Melissa Luckow (BH) greatly im-
proved the presentation of the paper.

744 Stipa 18(3)
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et th: C.R. Metcalfe & L. Chalk (editors). Anatomy of the Dictyledons. Vol. 1. Syshootatic
anatomy of leaf and shoot, with a brief history of the subject. Clarendon Press. Oxtord, U.K
Lavin, M. & M. Luckow. 1993. Origins and relationships of — North America in che
context of the Boreotropics Hypothesis. Amer. J. Bot. 80:1-14
Linpey, J. 1848. Hluscraced dictionary of
nary of botanical terms by John Lindley. [Reprint, With an Introduccion by Alice Eastwood,

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eal

ysotanical terms. Excerpt

Stanford rans rsity, School of Earth Sciences, 1964}.
LuNbeLL, C.L. 1966. Myrsinaceae. In: P.C. Standley & L.O. Williams, eds. Flora of Guate-
mala, Part a Number |. Fieldiana, Bot. 24 (8/2):135—200
ee Flora of Panama, Part VIII. Family 150. Myrsinaceae. Ann. Mis-
sourt Bot. Card. 285-353
a Neotropical Myrsinaceae—VI. Phytologia 49:341—354.
: —. 1982. Neotropical Myrsinaceae—VII. Wrightia 7:38-50.

Mercare, CLR. 984 . Some basic types of cells and tissues. Pp. 54-62. In: C.R. Metcalfe
and L. Chalk, eds. Anatomy of che Dictyledons. Vol. |. Syshootatic anatomy of leaf and
shoor, with a brief history of the subject. Clarendon Press. Oxford, U.K

Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9):1-437.

Nayar, M.P.& GS. Girt. 1975 [1976]. A synopsis of the genus Hymenandra A. DC. (Myrsinaceae)
and a new ae from Burma. J. Bom. Nat. Hist. Soc. 72:818-821.

Pipoty, J. 1987. A systematic revision of the genus C a mithus subgenus Grammadenia
(Myrsinaceae). erate New York Bot. Gard. 43:1—7

_ 1992a. The ae hi subgenus C re
Ann. Missin — a 79:90
9 2b. Ardista ne (Myrsinaceae): A new species from the Antioquian

ne

rudayana.

Chocé of C — Novon 2:389—39 1.

Pipoiy, J. AND J. RICKETSON. 1998. A revision of the genus Ardisia subgenus Graphariisia
reiemE gts Sida 18:435—472.

RICKETSON, J. AND J. Cie, 1997. A. synopsis of the genus Gentlea (Myrsinaceae) and a
ey t - genera of Myrsinaceae in Mesoamericana . Sida 17:697—707

STAHL, 1996. The relationships of Heberdenta bahamensis and H. pen Dilip 7 (Myrsinaceac).

Bor. J. Linn. Soc 122:315-334

PIPOLY AND RICKETSON, Hymenandra and its boreotropical implications 745

STANDLEY, 1938. Flora of Costa Rica: Myrsinaceae. Publ. Field Mus. Nat. Hist., Bot.
Ser. 18: vies 0.

STONE, B.C. 1991 {1992}. New anc eee rears Myrsinaceae, VI. Revision of the
genus Hymenandra A. DC. Gard. Bull. Sing. 43:1-

TirrNey, B. 1985a. Perspectives on ae origin ee the dosed similarity between eastern

Asia and eastern North America. J. Arn . 66:73-94,
ee 1985b. The Eocene Noth jane: land bridge: its importance in Ter-
tary aie ciedemn phytogeography of the northern hemisphere. J. Arn. Arb. 66:243—273.
oe b, W.L., J. KRAHULIK & R.C. RoLins. 1984. Trichome description and classification.
40-53. In: C.R. Metcalfe & L. Chalk, a Anatomy of the Dictyledons. Vol. 1. Syshootatic
anatomy of leafand shoot, with a brief history of the subject. Clarendon Press. Oxford, U.K.
WENDT, T. 1988. Chiangiodendr on (Flacourtiaceae: Pangieae), a new genus from southeast-
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—___. 1989. Las selvas de Uxpanapa, Veracruz-Oaxaca, Mexico: eae de
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oo S. 1990. A monograph of Sabal (Arecaceae: Coryphoideae). Aliso 12:583-666.

NUMERICAL LIST OF HYMENANDRA TAXA

H. stenophyHa (Donn. Sm.) Pipoly & Ricketson
H. calycosa (Hemsl.) Pipoly & Ricketson

H. sordida (Lundell) Pipoly & Ricketson

H. squamata (Lundell) Pipoly & Ricketson

_H. wilburiana (Lundell) Pipoly & Ricketson

_ H. pittieri (Mez) Pipoly & Ricketson

_H. callejasti (Pipoly) Pipoly & Ricketson

_ AL. acntissima (Cuatrec.) Pipoly & Ricketson

_ H. crosbyi (Lundell) Pipoly & Ricketson

O AOR ee Doe

wo ~

b,

LIST OF EXSICCATAE
Figures in parentheses refer to the numbers from the numerical list of taxa.
Collection numbers in boldface type indicate type specimens.
Aguilar, R., et al., 3663 (6); Alfaro, E., 169 (2); Allen, P., 2056 (6); 2176 (6); 3806 (6): 5828 (G):
S828A (6); 6726 (6); Angulo, L., 394 (6); Antonio, T., 1364 (6); 3627 (6); 3632 (6): Argiiello M., D.
et al., 11 (6).

Baker, R. & W. Burger, 126 (6); Bello C., E., 1059 (3); Brenes, A., pass 5 (2); 21243 (2); Burger, W.
& R. Baker, 9869 (2); Burger, W. & G. U., 4406 (6): 4644 ( ); Burger, W. & R. Stolze, 3738
(6); 5853 (4); Burger, W. et al., 11687 (2)

Callejas, R. et al., 5683 (7); Carvajal U., A., 392 (2); Chacén, A., 934 (G6); Chacén, I., 80 (5); 717 (5);
ae, I. & G. Herrera, 1758 (6); Chive, C., 71 (3); Chinchilla, M., 93 (2); 100 (2); Churchill, H.,

3867 (6): 3876 (6): ee as Gs 12 1): 3:70 (1): Croat, T., 36246 (2); 36703 (6); 43538 (3); 44581
(G); Cirtredasas, J. 7219 (8); 261 2 (7

746 Sipa 18(3)
W. & J. D'Arcy, 6749 (6); D'Arcy, W. & B. Hammel, 12292 (6); Davidse, G. & C. He sie
age Davidse, G. & G. Herrera, 31379 (5); Dressler, R., 4254 (6); 5962 (6); Dressler, R. &

25 (6); Duke, J., 12197 (6); Dwyer, J., 835 1 (6); 8551 (6); 11869 (6); Dwyer, J. & B. Lee
ee ‘6 8690A (6); Dwyer, J. et al.,4476 (6).
Folsom, J., 1242 (6); 3163 (6); ree a . R. Page, 5970 (9); Folsom, J. et al., 5741 (G); Fonnegra
et al., 953 (7); 1228 (7); 1249 (7); 13
Garcia, D., 112 = 312 (2); Garwood, N. et al., 936 (5), eee A., 2011 (2); 6854 (6); me (6);

Gentry, A. et al 3939 (2); Gomez, L. et al., 20415 (1); 23383 (4); Gémez-Laurito, - 8 (1);
srl M., 237] os Grayum, M.& R. epee (4); . M. &G. Her 2 (2);

857 (A); ): 9200 (6G); Grayum, M. & B. os 53 Gri ae M. et al., 3369 (6); ete 4969
Grijalva, A. & D. Bradford, 3652 (2); nae . tal., 3393 (2).

Haber, W. & E. Bello C., 1714 (3); 7433 (2); Haber, W. et al., 4430 (2); 4432 (2); 4447 (2); | 2 (2);
4759 (2): 4848 (2); Hammel, B., 958 (6); 1928 (6); 2662 er 3220 (6): an (6); 4054 (6); pe 3 (A );
8762 (4); 10962 (4); 2372 (4); 13717 (1); Hammel, B. & M. Chavarria, 17073 (4); Hammel, B. & M.
Grayum 14342 (1); Hammel, B. & J. Trainer, 12757 (4); 13262 (5); Hammel, B. et al., 14078 (3);
15186 (6); 17370 (2); 17829 (5); Herrera, G., 6279 (3); Herrera, H. & P. Perez, 98 (6).

ee
—

Jacobs, B., 2377 (4); 2917 (5); 3113 (4); 3189 (5); Jiménez, A., 3738 (6); Jiménez, Q. et al., 709 (2

Khan et al., F 1141 (2); Knapp, $., 1965 (6).

anes B.,35A es - Liesner, R., 53066 (2); Liesner, R. et al., 15041 (2); 15108 (3); 15348 (6); Lobo,
7 (2), Lciselle, B., 144 (4); Luteyn, J. & H. Kennedy, 1704 (6).

Marin, J. € , 257 (6); ae 3., 2250 (6); McPherson, G., 11270 (6); Molina Be A. et al.,
17525 (3); ies 190 (6); Moraga, C. 4 (2); Morales, J., 4354 (6); a 05 (2); Morales, J. & Y. Salazar,
195 (6); Morales, J. & V. Urena HON Morales, J. et al., 2544 (2); Moreno, A., 7 (1); Moreno, P.,
7532 (2); (2); 26053 (2); 26753 (2); Moreno, P. & J. Sandino, 12853 (2); Mori, S. . M. Crosby, 6313

(6); 6423 (9); Mori, S. & J. Kallunki, 2978 (6)

Nee, M. & B. Hansen, 14099 (1 a Newt D.& P. Vincelli, 3506 (5); vers, G. de, 7203 (6); Nevers,
G. de & H. Herrera, 4490 (9); 1 (9): Nevers, G. de et al., 3941 A 6398 (6); 6757 (9).

Opler, P., 188 (4); 332 (4); 380 (2),

Paredes, R., 944 (G6); Picado, A. ec al., 51 : Pictie _& A. Tonduz, 9173 (1); Poredat, L. et al.,
3681 (2): Porter, D. et al., 4422 (6); soe ens = 106 (6).

Quesada, F., 320 (6).

Ramirez, V. et al., 62 (3); Raven, iS 21829 (6); Rios, P. 374 (3); Robles, R., 1571 (5); 1666 (5);
badger, A. &N, Zamora, 2067

Sanchez, J. et al., 308 (5); Sandino, J., 159 (2), Schatz, G. & M. Grayum, 653 (5); 1303 Se 1) Sechube rt,
B., 1079 (2); Seibert, R., 1572 (1); Smith, A., 1630 (2); Sperry, J. 790 (4); 1726 (2); 2609 (2);

‘<

P& Hi Nee pa {4588 (2); 45539 (2); 45545 (2) ): 45862 (2); 46181 (2); 46221 (2); 47122 (6); ie
W., 23977 (5); Stevens, W. & O. ee 1432 (5); Stevens, W. et al., 24972 (5); Sullivan, G., 553
(6); eae K.& W. D'Arcy, 3565 (6) tsma, K. et al., 2453 e 4220 (9); 4279 (9).

Tate, R., 239 [454] (2); Taylor, C., 4551 (2); Todzia, C., 1502 (4); Taylor, C. & C. Skotak 4630 (3);
Tonduz, A., 9586 (1); 13369 (6).

Utley, J. & K., 5318 (2)
Valerio, J., 147 (2); Villalobos, R., 37 (2).

Wilbur, R., ey a 40442 (A); 64246 (4); Wilbur, R. & B. Jacobs 34720 (4); 34727 (4); Wilbur,
R. & D. Stone, 9745 '

Zamora, N. & Q. Jiménez, 1268 (6).

MYRSINE LUAE (MYRSINACEAE),
A NEW SPECIES FROM BRAZIL
JON M. RICKETSON
Muassourit Botanical Garden
P.O. Box 299, St. Louis MO 63166-0299, U.S.A.
jon. ricketson@mobot.org; jricketson@lehmann. mobot.org
JOHN J. PIPOLY II

Botanical Research Institute of Texas
509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.
Jpipoly @ brit.org; clusia@latinmail.com

ABSTRACT
A review of peter specimens of Myrsine efor - Flora anneNe Veneenage oe dd project
revealed the Pp new taxon Serra de Lua erritorio do
Roraima, Brazil. Ene is described and ne and its relationship to its nearest

gener, Myrsine guzanensis is discussed. An outline map showing the type ee is ser ie

RESUMEN

Al estudiar pliegos del herbario pertenecientes al género Myrsine ees la guayana venezolana,
se encontré una especie nueva para la ciencia proveniente de la Serra de Lua, del Territorio
do Roraima, Brazil; la zona brasilefia adyacente a la guayana — Se describe, se
ilustre y se discute el parentezco de la nueva especie, Myrsine luae. También incluye un mapa

mostrando la localidad tipa.
INTRODUCTION

The genus Myrsze R. Br. contains ca. 300 species of which nearly 1/4
remain undescribed. C. Chen and Pipoly (1996), Pipoly (1991, 1992a, 1992b,
1996), Pipoly and C. Chen (1995), and Ricketson and Pipoly (1997) have
provided summaries of evidence for broader circumscription of the genus,
especially to include Rapanea Aubl. Myrsine is pantropically distributed, occurring
in diverse vegetation types, from mangroves to subalpine scrub, but always
in moist, wet or pluvial habitats. The genus is defined by lateral (axillary),
fasiculate or umbellate inflorescences, sessile or on short, perennating pe-
duncles girdled by persistent floral bracts, thus forming “short shoots.” In
preparation for our treatment of the genus Myrsine for Flora of the Venezuelian
Guyana, a new species from just out side the region in the state of Roraima
in Brazil was discovered and is described herewith.

Myrsine luae Ricketson & Pipoly, — ae (Fig. 1). Typr. BRAZIL. Rorama:

Summit of Serra da Lua, 02° 25—29' N ° 11-14' W, 1,400 m, 24 Jan 1969 (fr), G.
Prance, J. Steward, J. Ramos & L. Farias ie (HOLOTYPE: NY; isoryPes: BRIT, INPA n.v.).

Sipa 18(3): 747-750. 1999

748 Sipa 18(3)

' 0
6s Mt. Roraima
VEN ,
Bolivar GUYANA
Pp 4
—
Essequibo
T.F.
Roraima
Amazonas

A

Serra da Lua

Pico da Neblina

Amazonas

Fic. 1. Distribution of Afyrszve dae Ricketson & Pipoly.

ioad ramulos glabros, lamina coriacea nitidaque, petiolos marginatos, inflorescentiam
—8-floribus, M. guianensi valde arcte affinis sed ab ea ramulis angulatis (non teretibus),
necnon meine angulatis (nec cereus) Pe antiis Coriaceis s (non chartaceis), stigmatibus

sinuato-capitatus (non conicis), denique vel ol leis ( globosis),
praeclare distat.

Tree 4 m tall, LO cm DBH. Branchlets ridged, 0.5 cm in diameter, gla-
brous apically. Leaver alternate; blades coriaceous, elliptic to oblanceolate,
4—9.5 cm long, 1.6—3.5 cm wide, apically acute, basally acute, decurrent

on petiole, smooth, shiny and nited above, dull and inconspicuously black
punctate below, the midrib slightly impressed above, prominently elevated
below, the secondary nerves not prominent, |3—15 pairs, the margins en-
tire, flat to subrevolute, opaque, minutely glandular-ciliolate in bud, gla-
brous at maturity; petioles marginate, 0.5—1 cm long, glabrous. Stamnate
inflorescence unknown. Pistillate inflorescence a subsessile, 5—8-flowered um-
bel; peduncle obsolete to 3 mm long; floral bracts, deltate, 1—1.2 mm long,
1—1.2 mm wide, apically acute, early caducous, glabrous, the margins ciliolate;
pedicels angulate, 2-3 mm long, glabrous. Staminate flowers S-merous; ca-
lyx coriaceous, cotyliform, 1—1.5 mm long, the tube 0.3—0.5 mm long, the
lobes ovate, |—-1.2 mm long, 0.8—1 mm wide, apically acute, conspicuously
punctate and punctate-lineate, See the margins entire, glandular-ciliolate;
corolla chartaceous, subrotate, 1.8—2.1 mm long, the tube 0.5—0.6 mm long,
the lobes lanceolate, 1.3—1.5 mm lane. 0.7-0.9 mm wide, apically acute,
densely and prominently punctate and punctate-lineate, glabrous, the margins

RICKETSON AND PipoLy, A new species of Myrsine from Brazil 749

Pic. 2. Myrszne uae Ricketson & Pipoly. A. Branchlet. B. Infructescence, showing obovoid
fruic shape. C. Pistillate corolla, showing hastate antherode and marginal glandular gran-
ules. A-C, drawn from holotype.

750 Sipa 18(3)

entire, minutely glandular-ciliolate; staminodes inserted at junction of co-
rolla tube and lobe, the sterile anthers subsessile, oblong, 0.9—1 mm long,
(.1—0.2 mm wide at apex, 0.4—0.5 mm wide at base, apically apiculate,
the apiculum proximally curved, basally deeply cordate, the connective epunctate;
pistil 1-1.9 mm long, the ovary globose, 1—1.2 mm long, 0.7—0.9 mm in
diameter, prominently pellucid punctate and punctate-lineate, the stigma
sessile, sinuate-capitate, prominently vertically lobed, 0.5—0.7 mm long,
0.5—0.8 mm wide, persistent in fruit, placenta globose, 2-ovulate. Fruit green,
cylindrical to obovate, 3.5—4 mm long, 2.4—3 mm in diameter at the apex,
1.2—1.7 mm in diameter at the base, prominently pellucid punctate and
punctate-lineate, glabrous.

Distribution. —Known only from the type on the summit of Serra da Lua,
Roraima, Brazil (Fig. 2), growing at 1,400 m elevation.

Ecology and conservation status. —Myrsine luae inhabits cloud forests in Serra
da Lua, a somewhat isolated range of mountains. No ecological notes are on
the type collection, but it is from an area known for granitic mountains
inhabited by vine forests, with cloud forests above.

Etymology.—Names for the location of the type locality, Serra da Lua, Territorio
do Roraima, Brazil.

Myrsine /nae is most closely related to M. guianensis, but 1s easily recog-
nized by its angulate branchlets and pedicels, coriaceous calyx sessile, sinu-
ate-Capitate stigma and ellipsoid or obovoid fruit.

ACKNOWLEDGMENTS
We thank the cited herbaria for loans of specimens, K. Gandhi and an
anonymous reviewer for critical reviews of the manuscript.
REFERENCES

C. Cuen, C. and J. Prrory. 1996. Myrsinaceae. In: Wu Zheng-yi and P. Raven, eds. Flora of
China. ey Press, Beijing and Missouri Botanical Garden, St. Louis, MO. 15:34—38.
Merz, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9): 1-437.
Pipoty, J. 1991. Systematic studies in the genus Myrsime L. (Myrsinaceae) in Guayana. Novon
204-210

199 2a. Estudios en el género Myrsine (Myrsinaceae) in Colombia. Caldasia

_ 1992b. N I Myrszne (Myrsi yin Peru. Novon 2:392—407.
1996. C eaeibubions at a new flora of the Philippines: I. A synop-
sis of the genus Myr (Myrsinaceae). Sida 17:115-162

ae C.C. CHEN. 1995. ee notes on the Myrsinaceae of China.
“Maven a: 357-3 61

RICKETSON, J. AND J. Piro 1997. Nomenclatural notes and a synopsis of the genus Myrsine
(Myrsinaceae) in Mesoamerica. Sida 17:579-589

BOTANICAL RESULTS FROM THE 1995
BISMARCK-RAMU EXPEDITION
IN PAPUA NEW GUINEA

W. TAKEUCHI

Botanical Research Institute of Texas
clo Papua New Guinea Forest Research Institute
and Lae National Herbarium
P.O. Box 314, Lae, PAPUA NEW GUINEA

ABSTRACT

Botanical results are presented from the 1995 biological survey of the Bismarck-Ramu
tract, an area identified by multiagency assessment as a prime site for possible conserva-
tion action. The findings provide confirmation of the area’s presumed biodiversity value.

er 610 distinct morphospecies and up to 15 confirmed or suspected plant novelties were
rere a an IREEOSING) 24-day ex pedition Taree’ ws aie new Species are re formally de-
scribed. General descrip f the

explored by the survey team. Adoption of s} ore management sad conservation measures

is recommended for the subject territory.

ABSTRACT (MELANESIAN TOK PIsIN)

Wanpela wok bus painimaut bilong ol saveman bilong ol bus diwai, rop, na gras, ibin
kamap long Bismak - Ramu territori long Oktoba 2 igo 24, 1995. Ol saveman bilong bus
ol i kaunim olsem moa long 610 ol kain kain diwai, rop, na gras i kamap long dispela hap.
Na wok painimaut i kamapim olsem fiftinpela long ol diwai na te i nupela kain olgeta.
Dispela tupela ten na foa de wok i soim dispela ples Bismak - Ramu em i holim wanpela
long ol kain ples bus we Papua New Guinea mas lukautim gut. ae long dispela pepa 1
tok klia long olgeta samting mipela lukim long Bismak-Ramu bus.

INTRODUCTION

Papua New Guinea (PNG) is a well known center for biological ende-
mism and diversification. It ranks among the world’s most botanically di-
verse countries, with a species-level floristic content variously estimated as
high as 15,000—20,000 Johns 1993), 20,000+ (Womersley 1978), and to
ca. 25,000 (Mathew 1995). As one of its principal missions, the Depart-
ment of Environment and Conservation (DEC) is responsible for identify-
ing specific areas of conservation value within PNG. Based on several DEC
internal evaluations and the Conservation Needs Assessment Report (Beehler
1993), a 168,000 hectare tract between Mt. Wilhelm and the Ramu River
was recently selected for potential designation as a conservation unit (Figs.
1-2). This poorly-known area of interest (AOI) was suspected of sequester-

Sipa 18(3): 751-782. 1999

Sipa 18(3)

solomon Sea

Coral Sea

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ee eee — oo ae tore ae an meer” 5 ies sa a = ya
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: é an ( | 44 Conservation AO! | ?
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Fic. |. Top. Papua New Guinea in aspect, showing the general location of the survey tract

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 753

on

ing high biodiversity, and a survey was deemed necessary to secure more
information on its suitability for further development as a protected zone.

During October 2—26, 1995, a biological survey was thus conducted from
4 base camps established between 600-2400 meters elevation inside the
AOI (Fig. 3). Ten scientific participants from Papua New Guinea evalu-
ated the botany, entomology, and vertebrate zoology of the area. Survey protocols
were consistent with the ICAD (Integrated Conservation and Development)
model. Although a comprehensive account of the expedition was published
by Hedemark et al. (1997), only a cursory description of the vegetation was
included since the botanical specimens had not been examined. Recently
completed determinations, an amended site by site overview of the vegeta-
tion, synopsis of notable collections, and an improved species list (Appen-
dix |) can now be provided. A wide range of organizational, socio-cultural,
and physical site information is presented in Hedemark et al. (ibid, to which
the reader is referred), and will not be repeated here.

METHODS

The botanical component of the survey consisted entirely of opportunis-
tic general collecting, employing the ‘Kostermans method’ of field-pack-
ing duplicates in 70% surgical spirit for deferred processing. A complete
set of the Bismarck-Ramu plant collections has been deposited at Lae Na-
tional Herbarium (LAE). Principal recipients of duplicate sets are A, BRIT,
K, and L; residual sheets are dispersed in no particular sequence and are
likely to be at the institutions of family specialists.

In genera for which revisions are available, vouchers were keyed to spe-
cies and the result confirmed against annotated sheets at LAE. For unrevised
groups, exsiccatae were matched against authentically identified material
and/or original descriptions. A number of specialists were also consulted;
their identifications are provided with attribution on the attached species
list (Appendix 1).

In the following account, taxa are referenced by the collection number
on which the claim is based. The text has been expanded from Hedemark
et al. (1997: 41-44) by addition of taxonomic detail resulting from the new
determinations.

VEGETATION SUMMARIES
Camp |
Location: Mt. Oipu, GPS 05° 35.5' x 144° 47.3', elevation 2360 m
Life zone: montane
Forest type: elfin or low stature mossy cloud forest
Collections sequence: 10381-10625

Sipa 18(3)

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}00} UO PaMO}}O} B]NOY NK

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oa \ garg WES 2 wees 7
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ge? 2 = BY f

Hedemark et al. (1997).

from

f

Trip route of the expedition. Reproduced

D.

2

FIG

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 755

At 2360 m, the expedition’s highest camp was established in mossy ridgeline
forest exposed to severe wind shear and constant misting. The vegetation
at this site was generally stunted, taxonomically depauperate, and struc-
turally unstratified. Stocking densities tended to be comparatively high due
to the small size of most trees, with boles and crowns exhibiting the scrag-
gly form and poor development typical of cloud stands. Due to steep slopes
and everwet conditions, plant communities reflected the influence of fre-
quent landslips and exposure of new earth. Successional montane taxa were
prominent components of the vegetation on unstable terrain. A Saurania
complex with S. cf. capitulata (10596), S. ilicifolia (10424), S. cf. naumannit
(10539), and a possible Sawrauia sp. nov. (10447, 10570), was characteris-
tic of colonizing communities. Trimenia papuana (10393, 10402) and the
family Theaceae; including Eurya cf. esas (10394), E. tigang (10421,
10571), and Ternstroemta brittentana (10532, 10624), were also common in
secondary growth. The regenerating facies was particularly striking and distinctive
due to the towering inflorescences of Harmsiopanax ingens ssp. ingens (10514).

On the summit ridge beginning at ca. 2400 m, arborescent growth con-
sisted of a stunted monolayer less than 4 m in height. The principal woody
taxa were Olearia rufa (10598), Prunus pullei (10608), Helicia microphylla (10609),
and Ascarina philippinensis (10613). Acsmithia reticulata (10603) was com-
mon in sheltered ravines off the crest.

On buttress ridges below the summit, the canopy layer was similarly reduced
to dwarfed scrub. Phanerophytes typically included Di/lenia cf. schlechteri/
quercifolia (10495), Elaeocarpus tartensts (10422, 10591), Garcinia archboldiana
(10494), Planchonella monticola (10589), Podocarpus sp. (10585, sterile),
Schuurmansia henningsii (represented by two distinct architectural morphs:
10515 robust arborescent, and 1/0409 dwarfed form), Weinmannia sp. (all
sightings sterile), and Xanthomyrtus montivaga (10602). Two Pandanus spe-
cies; one monocaulous and planted, the other branched and naturally re-
generating (§ Intraobtutus, 1/0623), were conspicuous emergents on sum-
mits and steep slopes, the latter species being otherwise scarce on level ground.
The understory was occupied by various infraspecific forms of the highly
variable Symplocos cochinchinensis (e.g., 10415, 10566, 10614). Schefflera shrubs
from the ‘S. schumanniana group’ were also common, with representatives
including S. schumanniana s. str. (10643, 10744), and allied forms such as
S. aff. sparsidentata 10427, 10471). Lianes were relatively rare and incon-
spicuous.

Canopy statures increased progressively on the path descending towards
Bubkile and lower slopes. However by elevation 2075 m, forest was replaced
by grassland composed of weed and waif species characteristic of anthropo-
genic disturbance. Eurya tigang (10571), Parasponia rigida (10567), Polyscias

756 Sipa 18(3)
belensis (1LO580), and Rhodomyrtus novoguineensis (10568), formed a seral bor-
der between forest and grassland.

From the standpoint of floristic richness, herbaceous plants collectively
represented the most important elements in the summit forest. Orchids and
ferns were undoubtedly the most speciose groups. The genus Cyrtandra had
more morphospecies than any other dicot understory constituent. Urticaceae
was especially varied; E/atostema blechnoides (10450, 10478), E. mongiense (10535),
E. morobense (10433), and E. tridens (10475) being particularly common. Debregeasia
was also frequent, but the genus is unrevised and there are no available bi-
nomials for the Papuasian species.

Two new taxa were discovered on Mt. Oipu, including a Pélea sp. nov.
(10481; also 10559, 10740 from Camp 2), and Prinus sp. nov. (10588).

While the community composition was unarguably that of a well-pre-
served native forest, several cosmopolitan weeds have encroached along es-
tablished trails on ridgelines and buttress crests. Although adventive spe-
cies are often ignored in biological estimates of site value, such plants are
convenient indicators of the proximity and intensity of human activity. At
elevations above 2200 m, alien species were limited to Bidens pilosa var. minor
(10458), Ageratum conyzoides (10429), and an unidentified subshrub (10425,
10431) cultivated as a village ornamental. The introduced elements are benign
herbs and not community-invasive taxa like Piper aduncum of disturbed lowland
environments. A planted Pandanus (att. 2jilianettii) was seen only as scat-
tered individuals along footpaths, but is ocherwise naturally-occurring in
the Highlands region.

On Hammermaster and Saunder’s (1995) system of vegetation classifi-
cation, the Mt. Oipu communities are assigned to structural code ‘L,’ ap-
plied to lower montane forests (above LOOO m) having dense, small-crowned
canopies. Such forests change progressively in composition and stature ac-
cording to elevation, eventually grading into the high montane formations

(loc. cit.: 14). The expedition’s ground survey indicates that che Mt. O1pu
summit is structurally and taxonomically very close to subtype code ‘Ls,’
referring to very small-crowned forest where emergents (except Pandanus)

are generally absent.

The Mc. Oipu vegetation would also be regarded as a ‘lower montane
rain forest’ on the system of Grubb and Stevens (1985). However the ap-
ied

—

parent equivalence hides significant clistinctions in the way the term is app
by the different authors. The Hammermaster and Saunders classification is
derived primarily from stand-level characters determined from aerial sur-
ve of identifying merchantable forest. Grubb and Stevens

vey, with the es

employ h ition criteria ap plicable only by ground inspection (e.g.,
pede of buttresses, woody climbers, cauliflory, drip tips, etc.) and is

purely phytoecological in orientation. Irrespective of the distinctions, Mt.

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 757

Oipu is clearly a lower montane rain forest sensu Grubb and Stevens, and
of the ‘mixed forest’ type with no single dominant component.

Camp 2

Location: Mt. Gulno, GPS 05° 32.7! x 144° 47.8', elevation 2045 m

Life zone: montane

Forest type: medium stature mossy cloud forest

Collections sequence: 10647-10768; 10626-10646 from transit between camps

On Mt. Gulno, the canopy was similar to that from the first site. Many
taxa were present at both camps, though apparently differing in their fre-
quencies. Because of logistical concerns, collections were generally not re-
peated between camps for plants thought to be conspecific.

Kania eugentoides (10048) was a dominant tree species around Camp 2.
Other common trees included Actinodaphne tomentosa (10752), Ascarina subsessilis
(10760), Caldcluvia rufa (10673), Cryptocarya nothofagetorum (10652, 10680,
10728), and Sloanea brachystyla (10747). The most frequent shrubs were Acronychia
ledermannii (10717, 10750), Dysoxylum enantiophylum (10751), Fittingia sp.
(10661, 10745), Myrsine lencantha (10573, 10671), Pittosporum sinnata var.
renuivalve (10655, 10689), Steganthera cf. insculpta (10672), and S. ilicifolia
(10749). Woody genera previously dominant on Mt. Oipu (Weinmannia,
Dillenia, Garcinia, etc.), became less common at the second site. In general,
phanerophyte taxa seemed more similarly abundant, though this was diffi-
cult co assess properly due to difficulties in making collections. Expedition
climbers were rendered ineffective by rain, heavy misting, and thick bryo-
phyte growth on tree trunks. Since the camp site was less exposed to wind,

canopies generally exceeded 10 m, also making vouchers comparatively more
difficult to secure.

As for camp 1, understory and epiphytic plants were very diverse, with
ferns and herbs accounting for the eee of collections. Ericaceae was
well- pe a Devens movobeensis (10658), Rhododendron anagalliflorum
s. str. (10686), R. beye ] 2 (10685 ‘ and R. wrightianum var. insulare
(10656). The most sotabl le ae was a new species of Bulbophyllum (10724,
det. N.H.S. Howcroft).

During the hike between camps | and 2, tall stature forest was encoun-

tered along the descent to the Kanel River and on the corresponding climb up
opposing slopes to Mt. Gulno. A sharp structural break was evident at eleva-
tion 5700 feet aneroid (1740 m), with marked changes in epiphyte abun-
dance, forest stature, and tree architectural form. This elevational level probably
marks the lower limit of the cloud zone. The abruptness of physiognomic
transition suggests that the cloud line is spatially fixed, at least locally.

758 Sipa 18)

The Mt. Gulno vegetation is assignable to the same forest classification
units as discussed for Mt. Oipu.

Camp 3
Location: Wara Kanel (Kanel River), GPS 05° 31.8! x 144° 49.1', elevation
1545 m
Life zone: premontane
Forest type: tall stature ecotone forest with intermixed lowland and mon-
Cane taxa
Collections sequence: 10769-10964

Camp 3 was sited in secondary vegetation extending along the Kanel
riverbed and over adjacent slopes. Sewrauia and Cyathea were the most con-
spicuous members of this riverine formation. In the former genus, the ma-
jor species included Sauramia aff. conferta (10815), S. congestiflora (10826),
S. cf. naumannii (10904), S. schumanniana (10940), and an unusual subglabrous
species (10829, not S. plurilocularis). Cyathea angiensis (10848) and C. wernert
(10794) were common tree ferns in heliophytic situations, while C. hornes
(10846) was frequent in advanced regrowth. Gastonia spectabilis (10956) was
a massive emergent. Other plants indicative of disturbance were repeatedly
encountered throughout the area, their dominance the result of subsistence
agriculture by Gulno villagers. Euphorbiaceae and Piperaceae were promi-
nent in the successional vegetation, being represented by Euphorbia plumerioides
var. acuminata (10792), Homalanthus novo-guineensis (10957), Mallotus papuanus
(10947), Piper lessertianum (10874B, 10927), PB. radatzii (10773), and P. subbullatum
(10822). In mid-seral phases, A/phitonia excelsa sensu Schirarend (no coll.),
Melicope spp. (10823, 10892), Geunsia pentandra (10821), and Trema cannabina
(10918) became frequent, often forming dense stands. This regrowth asso-
ciation is found throughout northeastern PNG whenever human or natural
agencies permit establishment of seral taxa. Phytogeographically, the weedy
plants often range over the whole of Papuasia and also into the larger Malesian
region. Several alien species were noted, mainly ephemerals such as Crassocephalum
crepidioides (10799) and Erechtites valerianifolia (10797).

Away from the river, the primary forest canopy was diverse and more
difficult to characterize. Podocarpus (10952, sterile) and Calophyllum (popu-
lations sterile) were common trees in a premontane assemblage also com-

posed of Cunoniaceae, Elaeocarpaceae, Ericaceae, Lauraceae, Myrsinaceae,
and Winteraceae. Intermixing with montane taxa were lowland represen-
tatives from Moraceae (principally Ficws) and Meliaceae (Aglaia). Urticaceae,
Zingiberaceae, and various small ferns dominated the understory layer. Among
the more abundant gingers were A/pinia werneri (10869), Alpinia sp. §
Dieramalpinia (10964), Etlingera sp. (10878), Pleuranthodium sp. § Psychanthus

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 759

(10849, 10959), and Tapeinochilos (populations sterile). Common urticates
included Elatostema beccarii (10894), E. aff. belense (10891), E. novo-guineense
(10885, 10896), Poikilospermum inaequale (10858), and Procris frutescens (10901).
In general however, the forest floor community was sparse and taxonomi-
cally depauperate. The most notable collections were possible new species
in Dendrobium § Grastidium (108506; det. N.H.S. Howcroft) and Myrsine aff.
acrostica (10958; det. J. Pipoly).

Unlike the montane camps, epiphytic loads were minimal and tree boles
typically lacked mossy growth. Canopy species tended to be represented by
trees with massive stems branching high above the ground (>20 m). There
appeared to be sizable populations of merchantable E/aeocarpus and Syzygium,
but this could not be confirmed by fertile gatherings. In contrast to the
situation at the montane sites, virtually all the arborescent taxa in the ma-
ture growth forest were seen only in sterile condition. The few fertile indi-
viduals were often too high in the overstory for collection, the only excep-
tions being Ceratopetalum succirubrum (10853), Flindersia pimenteliana (10944),
and Lithocarpus cf. celebicus (10785). According to local respondents, the forest
trees begin flowering in November or December after the start of the rainy
season, a claim consistent with climatic data reviewed in Hedemark et al.
Dy):

The Kanal River forest is regarded as lower montane by Hammermaster
and Saunders (1995), but differs from the previous sites in the appearance
of lowland elements. Genera recorded from camp 3 which are characteris-
tic of the lowland zone include Caryota (sightings), Aceratinm (10806), Leea
(10825), Pometia (sightings), and Poikilospermum (10858). Lowland rain forest
can ascend to 1500 m (van Royen 1964), and the floristic composition of
the Kanal locality is transitional to such forests. Unlike high elevation for-
mations, the lowland-montane ecotone exemplified by the Kanal site has
not received critical attention in Papuasia (Grubb & Stevens 1985). Future
surveys in the Bismarck-Ramu tract could profitably focus on the transi-
tion, especially as unpublished findings from recent RAP (Rapid Assess-
ment Protocol) surveys suggest that floristic richness in Papuasia peaks at
or near this elevational level (e.g., Beehler 1997; Kulang et al. 1997).

Camp 4
Location: Wara Ikil (Ikil River), GPS 05° 30.8! x 144° 50.6', elevation 600

m
Life zone: lowland

Forest type: tall stature alluvial and foothill forest

Collections sequence: 10969-11110; 10965-10968 from transit between camps
4 to4

760 Sipa 18(3)

Flanked by steeply ascending ridges, camp 4 was the most isolated expe-
dition site and the least affected by disturbance. However on riverine flats,
the vegetation was still subject to natural upsets and marked by the devel-
opment of Pometia dominant canopy. Such communities are very typical of
lowland environments throughout northern PNG. In addition to Pometia
pinnata, the woody taxa on alluvial ground usually included Bridelia penangiana
var. penangiana (11020), Callicarpa longifolia (10973), Chisocheton lasiocarpus
(11000), Dolicholobinm oxylobum (1 1042), Endospermum labios (1 1048), Leucosyke
capitellata (11044), Mussaenda scratchleyi (11097), Pipturus argentens (10990),
Prunus dolichobotrys (1 1037), Saurania aft. conferta 1 1036), and Anacardiaceae.
The latter was represented primarily by sterile Bachanania, Campnosperma
brevipetiolata, and Semecarpus. Subarborescent Frevs was represented by large
populations of F. arbuscula (11038) and F. comitis (11100). The most com-
mon herbaceous plants were Derris cimeifolia (1 103 1), Desmodium sequax (1 1023),
Pueraria pulcherrima (1 1029), and Stachytarpheta cayennensis 1 1028). Urticaceae
was also common, with many sightings of Boehmeria platyphylla (110106),
Cypholophus nummularis (11018), Elatostema novo-guineense (11022), E. weinlandi
(11096), Laportea decumana (10974), and Poikilogyne macrophylla (11095).

The slopes and ridges above Wara Ikil have taxa less common than those
along the river and represent a more diverse forest. Proper botanical assess-
ment of such communities requires considerably more time and effort than
is possible with brief surveys such as ours. From general impression, the
stands near Camp 4 could be the richest plant community encountered by
the expedition. Both the Kanal and Ikil foothills would no doubt repay further
efforts at exploration.

Notable collections were A ntidesma katikit (11054, 11079), formerly known
only from a type collected in the Ramu area, Garcinia sp. nov. (1 1098, det.
P.E. Stevens), Psychotria sp. nov. (11090), and Syzygium sp. nov. (11068).

The Kanal River vegetation is assigned to type code’ Hm’ on Hammermaster
and Saunders (1995), a category consisting of medium crowned forests on
uplands below L000 m. The camp 4 area has the most merchantable tim-
ber seen during the survey and represents the forest type of greatest inter-
est to commercial operators.

NEW SPECIES, DISTRIBUTIONAL RECORDS, OR OTHER NOTEWORTHY COLLECTIONS

PTERIDOPHYTES

DENNSTAEDTIACEAE

Hypolepis scabristipes Brownsey; coll. 10778. Apparently a rare fern,
represented by few collections in the Malesian region (Brownsey 1987).
Hypolepis scabristipes is a distinctive species, with yellowish-brown sti-
pes marked by darker excrescences from the dilated hair bases (Loc. cit.).

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 761

THELYPTERIDACEAE

Sphaerostephanos sp. ?nov.; colls. 10707 and 10733. The genus is one of
the most speciose in Malesia (Holttum 1981) and includes numerous local-
ized endemics. Our expedition numbers are sessile-glandular on both lamina
surfaces, lack indusia, and have laxly setose sporangia. Sori are multiserial
and sometimes confluent. The collections are closest to S. adenostegius and
S. warburgit.

MONOCOTS

ORCHIDACEAE

Bulbophyllum sp. nov.; coll. 10724

Dendrobium sp., § Grastidium; possible sp. nov.; coll. 10856. About 45

orchid species were collected during the survey, from which orchidologist

N.H.S. Howcroft has determined two numbers as representing new species.
ZINGIBERACEAE

Alpinia sp. aff. odontonema, § Pycnanthus; coll. 10595. This differs from

A. odontonema s. str. in its sessile leaves with raised nervation on upper sur-

faces. The expedition’s collection conforms to material cited by R.M. Smith

(1978) as an undescribed species.

DICOTS

ARALIACEAE

Polyscias belensis Philipson; coll. 70580. An uncommon montane tree,
previously known from Bele River in West Iryan and from Morobe Prov-
ince (Philipson 1979). Lae Herbarium has only one sheet of this taxon. Apparently
a first record for the Highlands Provinces.

Resembles a Gastonia but the pedicels are distally articulated. The voucher
agrees in detail with the single sheet annotated by Philipson at LAE.
Schefflera aff. sparsidentata Frodin; colls. 10427, 10471. A possible novelty
in the S. ‘schumanniana-schraderiana complex’ from which a number of new
species have been described by Frodin (1982).

Closest to Schefflera sparsidentata but differing in the more robust inflo-
rescence with extended rachis, glabrescent axes, and much longer pedun-
culate umbellules. The flowers are distinctly pedicellate rather than sessile.
The conspicuous peduncular and floral bracts are densely clothed with setiform
innovations on margins and/or surfaces.

CLUSIACEAE/GUTTIFERAE

Garcinia sp. nov.; coll. 11098. A Garcinia with cordate-based leaves has

been preliminarily determined by P.F. Stevens as a novelty (pers. comm.).
The new species was seen as a single 10 m tree growing on the ridgeline

above camp 4. Its leaves are sessile, decussate, firm-coriaceous, and abaxially

762 Sipa 183)

glandular-lineate. It is vegetatively distinguishable from other species by
the subovate blade, typically 11.5 x 7 cm, with reflexed margins and
amplexicaulous cordate base. The flowers are deployed in numerous axil-
lary or infrafoliar fascicles, seemingly bisexual but probably functionally
unisexual and with the plants dioecious.

ERICACEAE

Rhododendron anagalliflorum Sleumer; colls. 10389, 10686. As delim-
ited by Craven (1980), R. anagalliflorum is an uncommon species confined
to the Carstenz Mts. and the Bismarck-Wahgi-Jimi Divide. It has the re-
duced leaves characteristic of Series Linnaeoidea.

The expedition vouchers were procumbent or decumbent epiphytes. Corollas
were campanulate, chartaceous, white or pink, with erect lobes and outer
surfaces exclusively lepidote. The ovary is densely clothed with patent hairs
but also provided with a lesser indumentum of coarsely tuberculate scales.
Styles did not exceed the ovary and were mostly glabrous.
EUPHORBIACEAE
Glochidion sp. nov.; coll. 11543. The collection was from a subarborescent
species seen in cloudy montane forest. It does not key out on Airy Shaw
(1980). In appearance most like Glochidion frodinii and G. urceolare, but separable
on the following combination of characters:

Vegetative parts hispidulous. Inflorescence axillary or internodal; pistil-
late flowers often solitary. Capsules globose, 1.5 cm diameter, subsessile,
glabrous, exocarp somewhat verrucose.

The capsules are eaten raw by Bubkile villagers, a practice not usually

encountered for Papuasian G/ochidion. The fruit is crunchy in consistency
and has a rather pleasant aftertaste. Unfortunately, this resulted in che vil-
lage laborers consuming all the gatherings as they were made, leaving only
a unicate for the press.
Macaranga reiteriana Pax & Hoffman; colls. 10496, 10508. Macaranga
reiteriana was formerly known only from Morobe Province, Gulf Province,
and the Idenburg River (Whitmore 1980). Lae Herbarium has material from
each of the areas cited in Whitmore (ibid) but no new occurrences have been
added to the national collection since then.

The species is ¢

on

istinguished by the single elongate stipule, narrow leaves,
and solitary fruits on bare peduncles. It is frequent in regrowth communi-
ties on Mt. Oipu. Recent work at Crater Mc. in Simbu (Chimbu) Province
has also documented the presence of the species from that area (e.g., Takeuchi
12262, 12274). Although the expedition vouchers represent a distributional
record for the High

—

ands region, the plant is almost certainly more com-
mon and widespread than herbarium specimens would indicate.
Mallotus papuanus (J.J. Sm.) Pax & Hoffman, or aff.; coll. 10947. The

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 763

species has paired leaves; each leaf pair consisting of a highly reduced, stipuliform
lamina opposed to an unreduced caudate blade. An indumentum of fulvous
hairs covers the apical parts and underleaves.

Matlotus papuanus was previously regarded as endemic to West Iryan (Airy
Shaw 1980). The expedition collection is apparently a first record for Papua
New Guinea. Although annotated specimens of M. papuanus have not been
seen, the species’ characteristics are sufficiently distinctive for a descrip-
tion-based identification.

EUPHORBIACEAE/STILAGINACEAE
Antidesma aff. chalaranthum Airy Shaw; coll. 10716 (fr). Antidesma
chalaranthum is known with certainty only from the staminate type collec-
tion, obtained from Goroka subdistrict in the Eastern Highlands (Airy Shaw
1979). A second specimen (Stres¢mann & Kairo NGF 27636), was referred to
this species as an example of the female plant, though the assignment was
explicitly provisional (loc. cit.). The expedition voucher is similar to NGF
27630; both numbers being subappressedly puberulent on twigs, inflores-
cence axes, and abaxial midveins. Fruits are also identically glabrous, 5 mm
in diameter, and with lateral styles. However our Bismarck collection has
drupes distinctly oblique, compressed, and lacking a thin-crustaceous pericarp;
characters unlike the number cited by Airy Shaw. There are possibly two
taxa hidden in the chalaranthum facies.
Antidesma katikii Airy Shaw; colls. 1/054, 11079. Supposedly a rare endemic,
previously known only from the type specimen (Coode & Katitk NGF 32762)
originating near the Ramu River at 90 m elevation. The large linear-lan-
ceolate leaves with pubescent midrib readily identify the species (Airy Shaw
1973, 1980). Unlike the type, the expedition collections include flowering
material, from which the following accessory description is provided:
Inflorescence from leaf-bearing or defoliate nodes, axillary, racemose, 2—
6 cm long, rachis patently pubescent; bracts ovate, 0.5 mm long; pedicels
to 1 mm, provided with indumentum like the rachis; perigone cotyliform,
typically 1 mm Xx 0.5 mm, glabrescent or puberulous, margins minutely
toothed, otherwise truncate; disc tomentulose; ovary asymmetric, 0.8 mm
x 0.6 mm, pilosulous; styles excentric, 2—4, divergent or reflexed, 2-fid.
Antidesma katikiz is locally common and a characteristic taxon in the Kanal
drainage. The label on the type indicates that the species was very abun-
dant in disturbed forest at the original collection site. It is likely that the
plant is not as rare as the scarcity of specimens would suggest, but is sim-

ply undercollected and of limited range.

LAMIACEAE/LABIATAE

Basilicum sp.; coll. 10626. The collection is not B. polystachyon, the only
Basilicum species recorded for Papuasia (cf. Keng, 1978). It may represent a

764 Sipa 183)

new species or a distributional record. The plant’s major characteristics are:

Suffrutescent and terrestrial. Leaves opposite, herbaceous, sub-bullate,
adaxial surface dark green, abaxially purple. Inflorescence terminal only,
racemiform, axes puberulent; verticellasters about 1.5 cm apart, short and
sparingly branched; pedicels long. Calyx bilabiate, manifestly venose (also
with intercostals), hairs subulate and septate; lower lip with 3 segments,
midlobe furcate and biapiculate, lateral lobes much reduced, rounded or
obtuse; upper lip wider, entire, shorter than the lower labium. Corolla blue
to violet, bilabiate, tube contorted, shorter than the calyx; upper lip shal-
lowly 4-fid; lower lip induplicate, entire, enlarged, enfolding the stamens;
stamens didynamous, scarcely exserted, connate at the base, filaments gla-
brous but with some sort of median callosity, anthers discoid and centrifixed;
stigma 2-fid.

MELASTOMATACEAE

Astronidium novoguineense Merrill & Perry; coll. 10762. The species
was formerly reported only from West Iryan and is now newly recorded for
Papua New Guinea.

Collection 10762 keys out to couplets 71-72 in Maxwell & Veldkamp

(1990) and best matches A. novoguineense. However the shape of the calyx
tube also suggests A. fragilissimum. If the specimen 1s actually the latter
species, it would represent a first record for Mamose (i.e., northern PNG)
region, since A. fragilissimum is currently known only from Central Prov-
ince (oe, cit.)
Medinilla sp. nov.; coll. 10408. Keys to species 49-52 in Mansfeld (1925)
buc does not match the binomials there. The collection is somewhat like
Merrill and Perry’s ‘mansfeldiana-markgrafil’ group except for the cernuous
cauline inflorescence. Other salient characteristics are the following:

Epiphytic, erect, monocaulous or not. Stems and innovations setose. Leaves
opposite, acroscopically directed, elliptic, to ca. 20 cm X 7.5 cm, glabrous
except near the base of abaxial costae, S—7 plinerved. Petioles proximally
provided with large auriculiform alae, the auricles foliaceous, paired, purple,
rounded but with margins erose. Inflorescence racemiform or sparingly ramifying,
conspicuously and persistently bracteate.

MONIMIACEAE

Steganthera insculpta Perkins; colls. 10513, 10072. Steganthera insculpta
is the only species in Philipson’s (1986) conspectus with subsessile female
inflorescences. It was previously known from two collections in the Sepik
region (ibid).

MYRSINACEAE

Myrsine aff. acrostica (Mez) Pipoly; coll. 10958. Myrsine is currently be-

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 765

ing reviewed by J. Pipoly, and Malesian taxa formerly included in Rapanea
are being transferred to Myrsine.

Collection 10958 is either a new species or a very aberrant Myrsine acrostica
(Pipoly, pers. comm.).

MYRTACEAE

Syzygium sp., aff. megistophyllum Merrill & Perry; coll. 1/068. The collection
keys out to S. megistophyllum in Hartley and Perry (1973). It differs from
that species in the linear-elliptic leaves, to 36 cm X 8.5 cm in size, with
base subsessile-emarginate but not deeply cordate. The lateral veins are in
25-30 pairs, generally straight, and obliquely diverging to a commissural
nerve 2 mm from the leaf margin.

Syzygium megistophyllum is apparently known only from a fruiting speci-
men obtained in West Iryan (loc. cit.). The inflorescence on / 1068 was cauline,
developing as abbreviate cymes < 4 cm length at the base of a short stem.
This is not too different from the description for 8. megistophyllum, though
the foliar characters are otherwise distinct. The expedition voucher prob-
ably represents a new species.

PIPERACEAE

Piper lessertianum (Miq.) C. DC.; colls. /0874B and 10927. The species

is a laxly pubescent climber with auriculate leaves. It is infrequently col-

lected and apparently uncommon; Lae Herbarium has only two sheets of

this taxon.

ROSACEAE

Prunus gideonii Takeuchi, sp. nov. (Fig. 4). Tyeus: PAPUA NEW GUINEA.

ESTERN HIGHLANDS PROVINCE: ee Range, Mt. Oipu, ridge between ‘Camp
‘and Bubkile, GPS lat. 05° 35.513'S, x long. 144° 47.252' E, elevation 2357 m,
‘ Oct 1995 (fr), WY. Takeuchi ere (HOLOTYPE: LAE; tsorype: L).

AP. pulleo laminis | | lip | oblongis) 15 cm longioribus (nec minoribus),

atque plandularibus basalaribus sisolees an praeditis), denique habito monoaxiali (non
polyaxiali) facile dignoscenda.

Understory shrub, monocaulous or hardly branched at the top; provided
with fulvous to orange-brown indumentum on all vegetative parts. Stem
ascending, weak, virgate, 2-3 m long, woody throughout. Branchlets few
or none, if present short and obliquely ascending, tomentose. Stipules con-
spicuous, paired at the petiole base, free, lanate, persisting, acuminate, 1 2—
20 mm X 5—10 mm. Leaves spirally arranged, firm, manifestly bullate; mature
blades lanceolate to oblong-lanceolate, 18—27 cm x 9-11 cm, apex acute,
margin reflexed, base obtuse or subequally notched, often induplicated; adaxial
surfaces dark green, initially pilose on veins, later glabrescent, abaxially orange-
brown and lanate, opaquely punctulate; lateral veins 6—9 pairs, obliquely
diverging, supramedially looping and usually closing at 2-4 mm from the

Sipa 183)

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yt ile

re leaves. B: shoot with stipules

atut

4. Prunus |

Fic.

videonit Takeuchi, sp. nov. A: habit, m

leaves. C: infructescence.

and immature

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 767

margin, nervation deeply impressed above, on undersides very elevate, ar-
eolate and with blackened nerves; basal glands absent; petioles 12-15 mm
long, entirely pubescent. Inflorescence not seen. Infructescence racemose, soli-
tary, axillary from attached leaves, 4—-8.5 cm long, rachis 3-4 mm es
lanate; pedicels 4-12 mm long, densely pubescent. Drape ovoid to subg ;
12-13 mm X 9-10.5 mm, hypanthium residue 3—5 mm diam.; —
pinkish-red, appressedly hairy; style persisting, stigma capitate or some-
what discoid; endocarp glabrous. Seed single, abortive or immature, crispate;
testa glabrous.

vality,
in stunted montane forest within the cloud zone.

Etymology.—The new species is named for Dr. Osia Gideon, a specialist
in Papuasian Rubiaceae and Costaceae, and currently the deputy director
of the PNG Forest Research Institute.

Prunus gideonii has a distinctive aspect, with fulvous-lanate hairs on nearly
all parts and large bullate leaves to 25 cm X 11 cm. Other diagnostic fea-
tures are the sub-monocaulous habit and the conspicuous, persisting stipules.

The plant’s sectional affinity is unclear because the type is apparently
aglandular and lacks flowers. On the basis of phytogeography, P. gideonzi
probably belongs to subgenus Lawrocerasus section Mesopygeum (cf. Kalkman
1965). The type keys out to Prunus pullez in Kalkman (1993) but is obvi-
ously not that species.

On Kalkman’s (ibid: 322-26) key to fruiting specimens, the simplest
way of accommodating the new binomial is by deleting P. pw//ei from fork
46 but retaining line 46b as the lead to the following couplet:

Ramiform trees or shrubs; leaves elliptic to oblong, 2-12 cm length, basal
rlands present Prunus pullei
ieee or sparingly branched shrubs; leaves lanceolate, >15 cm length,
basal glands absent Prunus gideonii
RUBIACEAE
Psychotria howcroftiit Takeuchi, sp. nov. (Fig. 9). Typus: sears NEW GUINEA.
MADANG PROVINCE: Bismarck Range, ridge above ‘Camp

, Ge
x long. 144° 50.646' E, elevation 900 m, 23 Oct 1995 a fr), W. ioe 11090
(HOLOTYPE: LAE; isotypes: A, BISH, BRIT).
Propter inflorescentiam trichotomam, stipulas valvatas, tub rollir 2mm ee
denique fructum magnum, P. solomonensi vale arcte attints, age ap ea laminis 2 = a non

17-20) cm longis, nerviis secundariis 15
(non sessilibus) statim distinguitur.
Fruticose or subarborescent to 4 m height, vegetative parts entirely gla-
brous. Stem erect and laxly branching, basal swell absent. Branchlets terete,
4—7 mm diam., pithy, fleshy, moderately robust, collapsing when dry, sur-
faces smooth and nitid. Stipules valvate, basally connate, tfugacious, at first

768 Sipa 18(3)

Fic. 5. Psychotria howcroftit Takeuchi, sp. nov. A: habit; mature leaves and young inflores-
cence. B: architectural form (ternate) of the infructescence. C: drupe in longisection. D
and E: drupe in cross-section. Endosperm ruminations not shown.

acuminate and conduplicate, expanding and ovate when fully developed,

—9 mm X 6 mm, undulate, margins entire. Leaves opposite, coriaceous,
spreading, domatia lacking; mature lamina oblanceolate-oblong, 22—30 cm
x 8-13 cm, apex abruptly acuminate, base cuneate; adaxial surfaces dark

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 769

green, abaxially very pale green, in sicco bifacially fuscous; lateral veins in
16-19 pairs, equispaced, evenly arcuate, closing only rarely by marginal
loops, major veins embossed on upper side, prominulous beneath; reticu-
lum feeble, the crossing nerves subscalariform, otherwise with plexus ir-
regular; petioles 4—6 cm. Inflorescence terminal, to 5 cm length, umbelliform,
3 rachises connivent and approximately ‘trichotomous’, primary branches
ternate-verticellate, ultimately cymose, all axial surfaces light green and
glabrous; bracts caducous, deltoid, minute, reflexed, adaxially pilosulous.
Flowers sessile and externally glabrous; calyx cupular-turbinate, 5-6 mm x
4-6 mm, margins strictly truncate; corolla 4-merous, valvate, white, the
bud acute, 9 mm X 2.5 mm prior to anthesis, lobes oblong, divided to 3
mm from the base (rehydrated bud 12.5 mm x 4.5 mm with segments 7.5
mm X 3 mm and divided to 5 mm from the base); throat pilose, hairs sep-
tate-moniliform; stamens 4, alternipetalous, adnate near the sinuses; an-
thers dorsifixed, oblong; style exceeding the anthers, glabrous, ?heterostylous;
stigma 2-fid and fimbriate; disc annulate, glabrous, marked by a central
excavation after stylar abscission; open flowers not seen. Infructescence dif-
fusely paniculiform-umbelliform, flaccid, to 10 cm length, articulated at
the ramifications; peduncle 2 cm. Drupes subglobose-ellipsoid, 19—22 mm
(excluding calyx) X 15-17 mm, pericarp orange-red, convex, contracting
and conspicuously angulate after drying; calyx tube persisting at the sum-
mit, vasiform to cylindric, 3-4 mm X 4—6 mm. Pyrenes 2, equal, plane on
the commissural face, dorsally crested; endosperm ruminate.

Distribution and ecology—Known only from the type locality, there oc-
curring as scattered individuals in the understory of mature growth forest
or in stands with advanced regrowth.

Etymology.—The new species is named for N.H.S. Howcroft; an orchidologist,
silviculturist, and botanical illustrator, currently serving as the managing
consultant of a balsa project in New Britain.

Psychotria howcroftii is immediately distinguished by the exceptionally
large fruits borne ona diffuse, articulated infructescence. The oversized drupes
are made even more conspicuous by persistence of the 4 mm long calycine
tube. Among Papuasian Psychotria only P. monopedicellata has fruits as large but
that species has calyptrate stipules and monoaxial inflorescences (Sohmer 1988).

The immature inflorescence on P. howcroftii appears trichotomous (sensu
Sohmer), but since the mature infructescence is pedunculate, the initial
trichotomous structure is merely due to delayed prolongation of the pe-
duncle. Another developmental peculiarity is that stipular form becomes
manifest only at the subapical node, the stipules being otherwise severely
enfolded at the apex.

—y

770 Sipa 18(3)

The new species will key to fork 36 on Sohmer (1988: 15). It can then be
assimilated to the existing decision train by deleting lead 36b for P. solomonensis
and adding the following:

36b. Corolla tube at least 2 mm long
ateral veins >15 pairs; fruits >15 mm long....... P. howcroftii

Mature blades >20 cm long;
Mature blades <20 cm long; lateral veins <15 pairs; fruits smaller...... P. solomonensis
URTICACEAE
Pilea hedemarkii Takeuchi, sp. nov. Typus: PAPUA NEW GUINEA. Borper oF
WESTERN HIGHLANDS PROVINCE AND MADANG ProviNce: Bismarck Range, near ‘Camp
2’ on Mt. Gulno, GPS lat. 05° 32.7'S, x long. 144° 47.8! E, elevation 2040 m, 12
Oct 1995 (£1, fr), W. Takeuchi 10740 aoe LAE; tsorypes: K, L).

ao}

Species haec inter se allis species generibus, laminis linearibusque uninerviis atque squamibus
aurantiaco-lepidotibus indutis praeclare distat.

Weakly ascending monocauls or ramiform chamaephytes, <0.5 m height,
terrestrial, monoecious. S/ems slender, terete, glabrate, orange-brown, marked
by parallel cystoliths or not. St/pu/es axillary, connate, caducous, obscure,
0.2—0.3 mm long, entire. Leaves paired, isomorphous but generally unequal,
divergent, glabrous, chartaceous, bifacially squamulose; scales diffuse, peltately
based, orange-hyaline “a darkened centers; lamina linear, major blades
usually 17-34 mm Xx 2—3 mm, apex acute, margins distally and distantly
serrate, proximally entire, ‘base obtuse; adaxial surfaces dark green, provided
with linear cystoliths, these mostly transversal, less often randomly directed,
on leaf margins abruptly longitudinal and congested; abaxial surfaces pale
green to glaucous, collaterally glandular-lineate along the midrib; vena-
tion unicostate, other nervation invisible; petiole 1-4 mm long. Inflores-
cence axillary and solitary; cymes glomerulate, several together, occasionally
simple; peduncle obsolete or to 9 mm long and filiform. Ma/e flowers (rehy-
drated socal ect or less commonly pedicellate, bracteolate, glabrous,
entirely white; perigone 4-fid, typically 3 mm long, 1.5 mm wide at the
base, ovoid in bud, segments lanceolate-ovate, each about 1.8 mm xX 1.2
mm, costate, the rib excurrently corniculate or mucronulate; stamens 4,
oppositi-tepalous, adnate to the tube, filaments inflexed; pistillode reduced
toa minute flap. Female flowers glabrous, pedicelled or appearing sessile when
immature; perigone 3-partite, fleshy, lateral segments vestigial, median segment
enlarged and accrescent, acroscopic; staminodes 3, globular, oppositi-tepalous;
ovary ellipsoid to cylindriform at first, later oblique and compressed; stigma
penicillate, semi-persisting, directed at the major tepal. Infructescence en-
tirely light green. Fruits basiscopic ona retrorsely turned stalk, asymmetrically
ovoid, 1.2-1.0 mm Xx 1.0 mm, compressed, marginate; pericarp thin and
smooth. Seed flattened, rostrate.

Distribution and ecology. —Pilea hedemarkii is known from montane forest

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea Fes

in the Western Highlands, Madang, and West Sepik Provinces. During the
expedition it was often seen along footpaths and forest margins. Another
collection which is referable to this species (NGF 41691), has a label de-
scribing its habitat as ‘broken forest,’ further showing that the plant is found
in successional situations.

Etymology.—The new species is named for Michael Hedemark, the expe-
dition leader and a former conservation biologist with the United Nations
Development Program. He is currently with the Wildlife Conservation Society
in Laos.

Other Specimens Examined: PAPUA NEW GUINEA. West Sepik Province: Bli M
south of Oksapmin, broken forest on hillside, lat. 05° 20'S x 142° 15'E, elevation 7 00

2195 m), 22 Oct 1968 (fl), E. Henty, R. Isgar, G M. Galore, en (A, BRI, CANB,
K, L, LAE). Western Highlands Province: Bismarck Range, Mt. Oipu, subcrest slopes
of main ridge in vicinit ay of ‘Camp 1,’ low stature montane forest in cloud zone, GPS lat.
05° 35.5'S, * long. 144° 47.3'E, sivac on 2360 m, 05 Oct 1995 (fl), W. Takeuchi 10481
(A, BISH, BRIT, one L, LAE); Bismarck Range, Mt. ae ridge community between
‘Camp 1’ ane Bubkile, low stature montane forest in cloud zone, GPS lat. 05° 35.5'S

long. 144° 47.3' E, elevation 2400 m, 07 On 1995 (fl), W Takeuchi 10559 (K, LAE).

Papuasian Pilea have been treated in Winkler (1922) and van Royen (1982).
Although both authors provide keys to species «he coverage of either ac-
count is incomplete. The genus still awaits a synthetical revision.

Pilea hedemarkii is distinguished from all other Papuasian congeners by
the linear and uninerval leaves. The minute, orange-brown, and peltate scales
are also distinctive, though not unique.

DISCUSSION

The expedition discovered about 15 confirmed or suspected new species,
in addition to other noteworthy gatherings summarized in the preceding
section. At least 613 distinct morphospecies were represented in the 730
collections made by the survey. It is instructive to compare the number of
novelties reported here with two of the largest surveys recently concluded
from other parts of Papuasia, both of which were also of approximately one
month duration. The 1994 New Ireland survey produced only two new taxa
(Takeuchi & Pipoly 1998), and a total of 8 novelties is suspected from the
1997 Lakekamu survey (Takeuchi & Kulang1998). Results from the present
expedition support previous estimates of high biodiversity in the Bismarck-
Ramu tract, and justify enactment of conservation measures for the area.
Due to inclement weather and the overall scarcity of fertile sightings, the
collections coverage was far from comprehensive. There is clearly consider-
able scope for further discovery.

Sipa 183)

APPENDIX |

Expedition Plant List From Bismarck-Ramu. Numbers refer to exsiccatae

vouchers. SR =

sight record without exsiccatae; (m) =

male plant; (f) = fe-

male plant. Determinations by the author unless otherwise indicated. As-
terisk (*) preceding binomial indicates presence of discussion in text.

AVASCULAR PLANTS
BRYIDAE
genus indet.,
FERNS AND FERN ALLIES
ADIANTACEAE

Syngramma quinata (Hook.) Carruth., 11062
Taenitis blechnoides (Willd.) Swartz, 10937

10675, 11003

fOr EEN Acie?

Christ, nee 7, 10970, 11055

A seis affine Swartz, 105

Asplenium pit Bakes, sk ay ee
ude O9T¢

Aspleninm caudatum Vor

Asplenium morobense Copel., 10043B

Asplenium nidus L., SR from Camp

Asplenitum phyllitidus Don subsp. galas un
Holttum, /0893-A, 11050

Asplenium steerer Harrington, ree he 56

ei Harrington,

~
GN
x

ar

Aspleninm is at ginatum Rosenst.,
., SR from Camp 3

3

Aspleninm tenerum Forse.

= oo

Asplenium unilaterale Lam. 0

Dicdymochldena sles sata Saitek: 10924
ATHYRIACEAE

Diplazium bantamense Blume, 11064

Diplazinm cordifolium Blume, 10761, 11056

Diplazinm ais a tz.) Swartz, 10781

Diplazium sp.,

BLECHN ACEAE
iusculum (v.ANV.R.)C. Chr., 10435

Blechnum cf. archboldii C. ae 10604

Blechnum dentatum (Kuhn) Diels, LOBES, 11074

Blechnum deorso- -lobatum Brause,

irate ai: TOse6

air,

Stenochlaena areolaris (Harr.) Copel., 10772 2
CHEIROPLEURIACEAE
Cheiropleuria bicuspis (BL.) Presl, 10942

CYATHEACEAE
Cyathea angiensis (Gepp) Domin, eo 1S

Cyathea ct. cincinnata Brause, 107
Cyathea geluensis Rosenst., 10551, on 10706
Cyathea hornet (Baker) Copel., 10846

Cyathea perpelvigera v.AV.R., 1003 35, 10697
Cyathea wernert Rosenst., 1 0794
Dicksonia ct. sciurus C. Chr., 10765
DAVALLIACEA
Davallia divaricata Blume, 10776
Humata a , alpina-neoguineensis group’, 10572

, 11066
lene ee (Mett.) Copel., 10804, 10820
DENNSTAEDTIACEAE
Dennstaedtia glabrata (Cesati) C. Chr., 10688,
11087

Dennstaedtia sp., ee ao 10533
Histiopteris estipulata v , 10698

Hypolepis eer Brownses 10778
es sp., 1004
Paesta radula (Bake : C. Chr.,
DIPTERIDACEAE
Dipteris conjugata Reinw.,

10574

10617, 10803

GLEICHENIACEAE

I is (Burm.) Underw., SR from
este l
Gleichenia brassii C. Chr., 10611
Gleichenia dicarpa R. Br., 10616
GRAMMITIDACEAE

Calymmodon egelas = ) Copel., 10557
Ctenopteris sp. “Ci g *, 10575
Cc Balai Hogelliforme Brause, 0548

enopteris ct. longiceps (Rosenst.) C te 10594B,

10946
Crenopteris nied (Blume) i 10578
Crenopterts repandula Kunze,
Cc lal is elle . sae “10668

até NOL preri yetosa Cc Ip del., 1070

Cc tmp iene (Zoll .) Copel., ; 0549,
99

: Fae Pye

Copel.) Copel., /O594A,

10664
Grammtutis Cf. dolichosora (Copel.) Copel., 10915
10607
10594D
105406,

Grammitis interrupta (Baker) Copel.,
(Baker) Copel.,

(Baker) Copel.,

Grammitis scabristipes
Grammitis sumatranad
10594C

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea TID

Scleroglossum juncifolium Copel., 10627
Scleroglossum pusillum (Blume) v.A.v.R., 10390

HYMENOPHYLLACEAE

Hymenophyllum sensu lat
Mecodium sp., oni banlrinnn group’, 1/0679
aff. p

Mecodinm a Copel., 10839
Meringinm cf. ae (Copel.) Co oD 10621
cf. oe P 10550; sterile collection

ane u lat
Caplan bonsai Eres, 10916
Macroglena
Macroglena ree Baus) Cok, 10651
cf. Macroglena sp., 10721
Seis cf. sntermedta (v.d.B. ) Copel., 10969,
004

seh ae pallidum (Blume) Presl, 10628

Lindsaea group
Lindsaea obtusa J. Smith, 10691, 10844
Lindsaea obtusa J. Smith, 10705; pinnate form
Lindsaea pulchella (J. Smith) Mett. ex Kuhn,
10838, 10932
Lindsaea pulchella (J. Smith) Mect. ex Kuhn
var. blanda ae ex Kuhn) Kramer, 10525
Lindsaea rigida J. Smith, 10696
Sphoomers ee (L.) Maxon, SR from Camps
,&

Sphenomeris retusa (Cav.) Maxon, SR from Camps
1,3, &4
a sp., 10565

OMARIO CEAE
me trai ae) ee 11021
Ching, 10995,

Bolhitis rivularis (Brackenridge) Ching, 1/011,
large form

Sipe novoguineense Rosenst., 10590

Lomagramma sinuata C. Chr 998

EYCOPODIACESE

iifolia (Blume) Jermy, 10771
Rute eva i en mmaria (L.) Rothm., 1/05 Ey
Huperzia aff. ?pinifolia Trevisan, 108
Huperzia squarrosa (Forst. f.) Trevisan, "10935
Sie ites Forst. f., 97
L.) Vasc. & i ranco, SR from

Camps | & 3
MARATTIACEAE
Angiopteris evecta (Forst.) Hoffman, 10816
Marattia cf. tafaensis C. Chr., 10499, 10501
OLEANDRACEAE
Nephrolepis biserrata (Swartz) Schott, 10786
Nephrolepis hirsutula (Borst. f.) Presl, 11030

LV

Nephrolepis lauterbachit Christ, sen
Nephrolepts rosenstockit ee

Nephrolepis schlechteri Brause, nee 10769
Oleandra cuspidata Baker, - 4, ae 10775
Oleandra sibbaldii Grev.,

Oleandra werneri od ae

o

OPHIOGLOSSACEAE

Ophioglossum pendulum L., 10534

POLY PODIACEAE

Aglaomorpha drynarioides (Hook.) Roos, SR from
amp 3
A gl zomorph 1 heracle (Kunze) Copel., SR from

=

p3
Bet, mucronata (Kée) Copel. var. mucronata,
11017

Belvisia novoguineensis (Rosenst.) Copel., 10412,
10467, 10694

Belvisia validinervis (Kunze) Copel.

var. /i Bere Wal TW ] 2k

}5

1O50¢
Colysis polysora alas: C eine ae
es naria rigidula R from Camps

2&?
Goninon demersum (Brause) Rédl-Linder,
£0503
Goniophlebium pseudoconnatum (Copel.) Copel.,
10817

ah coe serratifolium Brackenridge, 10432,
10 0489
Lamp ees beg Baker, 1/0802,
distr. as L. curtisi1
Lenmapolom rd ‘sume Donk, 1/0783,
108,
ae een ne 10487
Microsorum papnanum (Baker) Parris, 10477, =
hymatosorus sp.
Microsorum sp., 10987, 11012
PI i Bl Pichi Sermolli,

ake

ume) Parris, 10505

10 782, 0871, sn, dices elie spp.
Be ee enervis (Cav.) Ching, 10528, 10759:
subgramineous,’ distr. as Crypsinus ee
slic hellwi ie cae Hovenkamp, 10449,

10558, 10931, distr. as Crypsinus senescens
Selliguea lauterba ie Bre Hovenkamp, 10439,
ibbs

>

istr. as i bbsia
Signs plat eae nridge, 10492; distr.
Sp.

PTERIDACEAE

Preris blumeana ae th, 10764
Preris wallichia ie Monel. 10493
Preris Big stc Christ, 10988

774
SCHIZAEACE
Schizaea peas (L.) J. eee
Schizaea fistulosa Labill., 107
SELAGINELLACEAE
Selaginella sp., angustiramea-hieronymidna group’ ,
dS

17, 10930

11057

nee

Tectaria group
iets cf. microthecia (Vée) C, Chr, & Ching,
1098
Tectarta ct. christovalensts
Tectaria decurrens (Presl) Copel.,
THELYPTERIDACEAE
Coryphopteris fasciculata (Fourn.) Holttum, 10620
Plesionenvon mavattiotdes (Alston) Holetum, 10547
Pneuniatopteris sogerensis (Gepp) Holttum, 10810
Pneumatopteris sp., ‘superba-subappendiculata grou
0490, 10545
Pronephrium beccarianum (Cesati) Holttum,

C. Chr.) Alston, / 1014
10997

som

0645
ae pentaphyllum (Rosenst.) Ho

ctum,
730
Prampian cf. scopvlorum Holttum, or aff.,
SA, 10780, LOSSO
ee ieee aurita (Hook.) Ching, 10485
Sp a a archboldit (C. Chr.) Holtcum,
)41

D3

Sphaerostephanos dimorphus (Brause) Holttum,
10788

Spl. ‘yf, BR Hol
10909

— unitus (L.) Holecum var. papilliferns
. Tolttum, 1/078

§ f on Holttum, /O808, 10971

10733

fo

‘poh sp. ?nov., "ri
f. Sphaerostephanos sp., 1080

VITTARIACEAE

Antrophyum plantaginenm (Cav.) Kaulfuss, 10542
Kaulfuss; s.l.,

Antrophyum reticnlatum (Korst.)
10898

10953
GYMNOSPERMS
GNETACEAE
Gnetum gnenon 1.., LOSOS

PODOCARI

sae imdb ‘catus (Blume) de Laub., 10899;

sterile specim
sas sp. A, ederm snti-nertiformis leaf type’,
sterile collection
oa sp. B, 10585; sterile collection

z var. angustifolia Holetum,

—

Sipa 18)

MONOCOTS

AGAVA

at

CEAE
Cordyline terminalis Kunth, SR from Camp 3

ARACEAE

Alocasia aequiloba N.E. Br., 11001
Alocasia nicolsonti A. Hay, 10497
4 f Ped pore aS fd [ sg | hore |

DI SR from
A
mp 4
Colocasia esculenta (LL amie eae
Cyrtosperma macrotum Becc. ex Engl.

ee amplissimum (Sc hott) E nals = on
mp 4

E ee pinnatum (1..)

ime a eee tee

omalonien

) Engl., SR

rc-stem herb
rdate-leat herb

10, robust ¢
110063,

7 Stone 's architecture

Homealomena sp i ! : :

Poth § Pathos,

ae sp., [0805
ARECACEAE/PALMAE

Calamus cf. reticulatus Becc., 10569

Caryota rumphiana Blume, SR from Camp 3

Heterospathe sp. A, 10706

Heterospathe sp. b 10676, 10954

Hydriastele sp., 11077

Korthalsia ct. a Blume, 1/061

Linospadix sp., 10712; not L. albertisiana

Orania ct. oreophila Essig, 10875

sterile collection

MMELINACEAI
Commelina diffusa Burm. f., 10789
Floscopa scandens Lour., 10873
Forrestia mollissima (Blume) Kds., 10989; as
Amischotolype £. marginata (Blume) Backer
Pollia thyrsiflora (Blume) Steud., 1 1076
COSTACEAE
Tapeinochilos hollrangii 1K. Schum., 10985; det
rideon
CYPERACEAE
Carex alopecnroides D. Don var. chlorostachys (D
Don) Clarke, 10455
¢ Boeck.,
ve , 10564
Cyperus cypermus (Retz. ol Ses 11086
Cyperus distans L. £., 10795
Cc nee aad E aoe 1070 10774
Sav.) Palla, 1/0629
10835

ened dichotoma (L.) V’ a

Hypoletrum compactum Nees & Mey., 10933

Hy poletram nemorum (Nahi) Spreng... 10900, 10912

— tea (Clarke) l Jictien, canal
serobiculata Nees & Mey., 1084

DIOSCOREACEAI
Dioscorea bulhifera \..,

11035

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea

FLAGELLARIACEAE
Flagellaria indica L., SR from Camp 3

HELICONIACEAE

Heliconia papuana WJ. Kress, 11045
JUNCACEAE

Juncus prismatocarpus R. Br. var. indicus, 10403
LILIACEAE

Dianella ensifolia L., 10419
MARAN E

TAC
Phrynium cf. ee ee K. Schum., 7/047
Phrynium sp., 10882
MUSACEAE
Musa sp., 11046
ORCHIDACEAE oc by N.H.S. Howcroft)

ae
si

Aglossorhyncha s
Bulbohby Mum "s a af eae 10384

Bulbophyllum sp., § Coelochilus, 10460

Bulbophyllum sp. 8 pa i oh 21

*Bulbophyllum sp. nov.,

Bulbophyllum sp., 10462, as 10949, 11091

Bulbophyllum or een sp., 10461

Cadetta aprina (J.J. Sm.) Schltr., 10387

Ceratostylis sp. 5

Chitonochilus Papuan Schltr., 10936

h

Aw

Coelogyne bec chb. 086 1
Coelogyne ne ite ae Ro Ife, O864
Dendrobium cuthbertsonii E. Vv. M. 386
Dendrobium lawesii Fv.) 095

Dendrobium cf. masarangense ie

10647

endrobium otaguroanum A.D. H
Dendrobinm aff. vexillarins J.J. Sm., 10459
Lum S Si ei 10586
Dag obium sp., § Eviopexts, 53
Dendrobium sp., hea teen possib

bp sp., 10472
a att. ramuana or neg 10633

oo. ot Eurycentrum, 10732; spur present
C

Glomera of aured eae 1063
Glomera sp., ie Be 10653
Glossorhyncha 7

aie a Sp

Lipari subs. enna cf. ne 10833
0

Liparis or Malaxis sp., 1092

a sien eu sp., 1/0391

Mediocalcar sp., 10610

Mischobulbum lancilabium Schlitr., 10939

Oberonia sp., LO836

Pedilochilus sp., 10669

Phreatia ct. petiolata Schletr., 10650
Phreatia sp., § Bulbophreatia, 10392
Phreatia sp., 10693

Platanthera papuana Schltr., 10692

ee

Podochilus na 105

Psenderia cf. pauci ae Schler., 2
Sti Hie Krzl., veee ae
Zen sp., § Hetaeropsis, 10560

PANDANACEAE

ees. 10640

€

Freycinetia cf. angustissima Ridley, 10725
Freycinetia sp. A, 10881
Freycinetia sp. B, s.n., Oct. 23, 1995
Pandanus sp., ‘adinobotrys- ayia group’, 10767
Pandanus sp., ass 1002.

Pandanus sp., 1107

a SIACEAE

yw A. Cunn., 10592B

iton ples sym ‘yh

POACEAE/GRAMIN
Bambusa cf. forbesii ily Holttum, 10962
: ae albens Trin.
ilbomarginat. mn 10448

sa myosotis Nee 10684

lsachne pauciflora Hack., 10938

Lophatherum gracile Brongn., 10847
Pennisetum macrostachyum (Bonen: ) Trin., 10798

Saccharum officinarum L., 10800
Setaria palmifolia (coani) Stapf, 10465

SMILACACEAE

Smilax cf. zeylanica L., 11034

ZINGIBERACEAE

Alpinia odontonema K. Schum., 10683

Alpinia werneri Valeton, 10869

Alpinia sp., + § Diramapinia, 10964

*Alpinia sp., a § Pycnanthus, 10595

linda id (Valeton DR. M. Smith, 10538

Etlingera sp. 8; Geanthus (Polyanthae)

ee ee sp. ¥? eda ites 10959

Riedelia geluensis (Laut.) V 597

Riedelia monticola ne ton, ies

Riedelia rosacea van Royen, or aff. man 7, 10708,
10757

Riedelia subulocalyx Valeton, 10723

DICOTS

f. 0 dontone CNA

CANTH

A ACEA
Hemigraphis aff. rina (Nees) E Vill., 77075

Leptosiphonium sp.
Pryssiglottis sp.,

Ol .
Rungia klossit 8. Moore, 10524

ACTINIDIACEAE

Saurania ct, capitulata A.C, Smith, or aff., 10596

es aff. pine Warburg, 10815, 11036
A.C. Smith, 1/0826

042 24

Saurania ilic cafe a van Ro en

Saurania y NaAHMaANNI Die or aff., 10539,
10904, 11043

Sauvania ‘crmaninue Diels, 10940

776

Saurania sp. A; sp. nov., 10447, 10570; no match
at LAE
Saurania sp. B, 10829; subglabrous

AMARANTHACEAE
Tresine herbstii Hook. f.,

ANACARDIACEAE
Campnosperma brevipetiolata Volk, SR from
Camp 4

10796

ANNONACEAE
Fd, hae Ib

11088
genus indet.,
APIACEAE
Hydrocotyle sibthorpioides Lamk, 10473, 10520
APOCYNACEAE
Alyxia markgraftt Tsiang, 11099
Parsonsia cf. sangitinea (Wernham) Markgr.,
Parsonsia sanguinea (Wernham) Markgt
(Markgr.) D.J. M I( 1735
Parsonsia warenensis Kanehira & Hatusima, 1/0906

10667

11093
r. var. brassit

Middleton,

Parsonsta sp., 108

bens pandacaqgui Lam., 10967

AQUIFOLIACEAE
Ilex scabridula Merrill & Perry, 10841
Ilex spicata Blume, 10583

ARALIACEAE
Gastonia spectabilis (Harms) Philipson, 10956
Harmsiopanax ingens Philipson ssp. sgens, LO5 14
Mackinlaya celebica (Harms) Philipson, 10674
Mackinlaya schlechteri Harms) Philipson, 10710
10968

Osmoxylon novoguineense (Schett.) Becc.,

*Polyscias belensis Philipson, 10580

efi schumanniana Harms ssp. schumanniana,
10643, 10744

*Sche fe esp. nov., af
10427, 10471

Sc hefflera “a straminea Frodin, 10498

Schefflera ct. versteegii Harms, 11109; also pos-
sibly 8. forbests

t. sparsidentata Frodin,

ARISTOLOCHIACEAE
10920; sterile collection

Aristolochia sp.,
ASCLEPIADACEAE
Hoya s

) p., 10922
T) lophora cissordes

Blume, / 1025

ASTERACEAE/COMPOSITAE
Adenostemma lavenia (L.)O. Krze., 10404, 10480
Ageratum conyzoides L., 10429

Arrhenechthites novoguinensis (S. Moore) Mattt.,

10482B ssp. novoguinensis

Bidens pilosa L. var. minor (Blume) Shertf, 10458

Blumea arfakiana acclle 10884

Sipa 18(3)
Blumea ices Mattt., - 8
Blumea sylvatica (Blume) DC.,
Blumea sylvatica lane De var, oniia

(Blu me) Randeria, a

Blume ca (Blume) D a atica, 10454
C Ae ORT Benth 38. ee la
Erechtites valertantfolia (Wolf) DC., 1
Mikania cordata (Burm. f.) B.L. ie Cae
— te Mattf. var. c/verea (Mattf.)

ae ra a ie 10598
Vernonia cuneata Less., 10831
BALSAMIN ACEAE
Impatiens hawkert Bull, 10381, 10428, 10872

BEGONIACEAE

Begonia cf. angustae Irmscher, 10992

Begonia sp., Raniensis group’, 10441; climber,
st. O. Gideon

Begonia tafe 10502,

afc
10890

ill & Perry, or aff.,

BIGNONIACEAE
Tecomanthe dendrophila (Blume) K. Schum. & Laut.,
10704

ieee ari EAE

“men tOS |

Lamk, SR from Camp 4

BURSERACE
beta ct. bas CX. Schum.) H.J. Lam,
5 ortbundus-versteeght?

mm
a

CAMPANULACEAE
Peracarpa ¢arnosa
Thompson, 10544

oo Hooker &

CARDIOPTERIDAC
Cardiopteris Sines onus 10793

CARYOPHYLLACEAE

gs ease oe Id. ex Roem. & Schult.,

SR from Camp .

CASUAR

pe ad Moore) L. Johnson, 10855
CELASTRACEAE

Celastrus monospermoides Loes., 10851
CHLORANTH

Ascii C B. Rob., /0613

Ascarina subsessilis Verdc., 10760
CLUSIACEAE/GUTTIFE

Garcinia archboldiana A. e a h, 10494

*Garcitia spn vf 1098, det. P.E. Stevens

Garcinia sp.,

C ae TE RONIACEAE

it (Plc inch. Pk inch. ex E ndl., 11104

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea

oe
Trichosanthes 0927A; sterile col
Zebneria eee (Jacobs) Jeffrey, 10737
Zehneria ct. cissymbium (Jacobs) Jeffrey, 10518

(a

ection

CUNONIACE
Acsmithta reticulata ae ) ne a Hoe 3
Caldcluvia rufa (Schler.) Hooglar 067
Ceratopetalum ee CT: ice 10853 3
PEPE NIP EN TEACEAY
le Gage var. gracile, 10584

DILLENIACEAE

Dillenia ct. schlechtert Diels or cf. quercifolia White
& Francis, 10495

ELAEOCARPACEAE
Aceratium parvifolinm Schler., 10806
Elaeocarpus tariensis Weibel, 10422, 10591
Sloanea brachystyla (Schietr.) A.C. Smith, 10747
Sloanea velutina (Schltr.) A.C. Smith, 10553

ELEAGNACEAE
Eleagnus triflora Roxb. cf. var. brevilimbata ‘T
10811, 10874

urt,

ICACEAE
Dimorphanthera cornuta J.J. Sm. var. tenuiflora

eumer, 10595
Dinette aff. cornuta J.J. Sm., 10743; det.

Dinmphte elegantissima IK. Schum. var.
ple vs (Sleu mer) P.F. Stevens, ese (fl);
det. Stevens

Dinsntedine se eee K. Schum. var.
splendens (Sleumer) P.F. Stevens, 11108 (fr)

Bree edulis sh, oe

0658
"adeno iste cee 10389,
0686

i lodendron beyerinckianum Koord., 10685

esac tron att. = erinckianum Koord., 10396;

the specie

ee ron ee Sleumer, or aff., 10863

Rhododendron cf. macgregoriae ar v.M. var. Ea
(J.J. Sm.) Sleumer, 1/07

Rhododendron superbum See 10859

Rhododendron wrightianum Koord. var. insulare
Sleumer, 10656

riers eae Sleumer, 10887

Vaccinium sp., § Oarianthe: aff. villosiflora mJ.J.Sm.,

10907

EUPHORBIACEAE
A aff. chalaranthum ie Shaw, eee
itidesma kattkii Airy Shaw, 11054, 11079
ae icaudata-squarrosa group’, ee
2 (m)

777

ches laxiflora Pax & Hoffman, 10582
Breynia cernua (Poir.) Muell. Arg., 10529
Bri ue bas Hook. f. ef. var. aes
110

nee coriaceo-lanatum ce Shaw, 1/0519
ae oan labios Schodde, 11048

uphorbia plumerioides Teijsm. ex Hassk. var.
_Sm., 10792

m

acuminata
sala Spe nov., 1054
F Muell., SR from Camp 4
— ses J.J. Sm., 10972
Macaranga candata Pax & Hoffman, 10414, 10470
* Macavanga reiteriana Pax & Hoffman, 10496,

+ Mavis ile (J.J. Sm.) Pax & Hoffman,
ff 47

Om eee v0 ; iy (Wark r) |
10407, 10957
FAGACEAE
Castanopsis acuminatissima (Blume) A. DC., 11082
Lithocarpus ct. celebicus (Mig.) Rehd., 10785
FLACOURTIACEAE
Flacourtia zippelii Slooten, 11039
Pangium edule Reinw., SR from Camp 4

Schum

GESNERICACEAE
Aeschynanthus sp. A, 10442
Aeschynanthus sp. B, 10630
Aeschynanthus sp. C, 10678
Cyrtandra fusco-vellea K. Schum., 11009
es udra aff. es aes Schler., 10897, 11078
Ze 43, 10741
wees nus Cyrtandra, 10510
eee on, 10469, 10715
Weare sp., cf. § Diplochiton, 10506, 10599
athe sp., cf. § Diplochiton or § Loxolobus,
1044€

Cyrtandra, 104.

Cyan sp., § Geodesme, 10383, 10452, 10681
Cyrtandra sp., a a ero 10577
0637, 108
Cyrtandra sp., a § Loxophyllum|Phaeotrichium,
Cyrtandra sp., § Macrocyrtandra, 10631
Cyrtandra sp., 10709; possibly = sp.10443
Dichrotrichum sp., 10636, :
Rhynchoglossum obliquum Blume, 10986
GOODENIACEAE
Scaevola oppositifolia R. Br., 10832

GROSSULARIACEAE
Polyosma aff. subalpina Schultz-Menz, 10555,
9677

HALORAGACEAE
Gonocarpus halconensis (Merrill) Orchard, 1OG00

778
Gunnera macrophylla Blume, 10420
ANGEACEAE
Dichroa febrifuga Lour., 104206; ‘sylvatica complex’
LAMIACEAE/LABIATAE
*Basilicum sp., 10626, 10972
Orthosiphon aristatus an ume) ie , 10966
Plectranthus parviflorus Willd., 81

LAURACEAE
Actinodaphne nitida Teschner, 11105
Actinodaphne tomentosa Teschner, 10752
10049
Cinnamomum ck. podagricum Kostermans, 10845

Cinnamomum clemensti Allen,

Oo
S
=
=
g
Q
i)
oh
=
ae
z
as
_.
~~
=)
Q
=
aw
és
ing
=)
5
EX
—
SS
a)
Ww"
~~
ON

Cryptocarya nothofagetorum Kostermans, 10652,
10680, 10728
Litsea carrvii Kostermans, 10444
LEEACEAE
Leea indica (Burm. ft.) Merrill,

OSAE/FABACEAE

10825, 11040

sequax Wall.
In al al rubidus ee re oerer group,
110 court

83; sensu
Mucuna novo-gnineensts Schett., SR from Camp 4
Pueraria pulcherrima (Koord.) yord.-Schumacher,

11029

LOGANIAC

EAE
Fagraea ct. cetlanica artis 10729
agraed reli ptica Roxb., 1110

Pagr
Geniostoma aft. ‘rupestre oe

LORANTHACEAE
Amyema squarrosum (Krause) Danser ssp.

10644

squarrosum, 10468
Macrosolen le (Lour.)
cochinchinensis, 1085:

Tiegh. var.

MAGNOLIACEAE
Elmerrillia tsiampaca (1..)
10738

Dandy ssp. tsrampaca,

MELASTOMATACEAE
Astronia atro-viridis Mansfeld, 10437, 10612
* A stronidinm ct. novoguineense Me on & Perry,
10702
Astronieae indet., 10523; probably Astronza sp.
Beccarianthus SPs 10902

Creochiton novoguineens! f.) Veldkamp &
Nayar, | 1094

Dissochaeta angiensis Ohwi, 10790

Medinilla albida Merrill & Perry, 10654

Medinilla aft. albida Merrill & Perry, 10556;

but leaves sessile

Sipa 1L8&(3)

Medinilla dentata Veldkamp, er
Medinilla ous ungiana Mansfeld,
ana-teysmannii group’, LOSO2;

852

Medinilla att. ‘lorentzi
to lorentziana

10406

Pe sogter1ensi3 oS or Pate

Medinilla ae Miq.,

gr.
Poikilogyne ie (Cogn. Sear 11095

MELIACEAE

Aglaia ct. silvestris (M. Roemer) Merrill, 10665;
but leaflets symmetric
— aff. ‘tomentosa group’, 10813; but inflo.

Ww

=)
(ay

yendant, ?
Cc pe heton lastocarpus ied )
formicarum-pachyrhat
eee 10438, 10751

Valeton, 11000;

MONIMIACEAE

Kibara karengana Philipson, 10554
c., 11049

Ramee ayacilisF aS. ona
Kanehira & Hatusima

va

1104 al
Steganthera ilicifolia A.C. Smith, 10593, 10749
*Steganthera insculpta Perkins, 10513, 10672

MORACEAE
Ficus adelpha Laut. & K. Schum.,
Ficus arbuscula Laut. & K. Schum.,
Ficus conttis King, ie aoe
Ficus opposita Mig., 107206,
ene — ex ee a from Camps

10903
11038

tie niin Diels, 10601
MYRISTICACEAE

Myristica pachyphylla A.C. Smith, 10660

Myristica subalulata Mig., 10787, 10929

Myristica velutina Markgr., 10711
MYRSINACEAE (dets. by J. Pipoly)

Ardista forbestt 8. Moore, or at. ‘ 1060

Ardista sp., 10718

Conand ey ee (Laut. & K.Schum.) Mez
0824

Mittingia sp., 10661, 1074

Maesa papuana oe eee: WT
Myrsine acrostica (Mez) Pipoly, 10
*Myrsine att. acrostica (Mez) Saat 10958;
Pca sp. nov
thea (IS Sc hum.) Pipoly, 10573, 1067

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea

MYRTA ;
Kania engentotdes Schler., 10540, 10648

Metrosideros eee Laut. var. humilis (Diels)

awson
Rhodomyrtus py eee Diels, /0568
Syzygium ck. longipes rill & Perry, 10914

ike malcense (L. ) Merrill & Perry, s.

aa . megistophyllum Merrill & Perry,
1068
Xanthomyrtus montivaga A.J. Scott, 10602

ENTHACEAE
Nepenthes maxima Nees, 10592, 10713
OCHNACEAE
Schuurmansia henningsit K. Schum., 10515;
characteristic robust for
hu

Schuurmansia henningsti K. Schum. , 10409; di-
minutive eon
PIPERACEAE
Peperomia ct. gurakorana Dull, 77067

Kunth, 10695
ee

io 10874B, 10927
975

Piper cf pndnbainene DC., or aff., 109
a K. Schum. & Laut., 10541, 10702,
1032

a ieee K. Schum. rae 10822

Piper subcanirameum C. DC.,

Piper triangulare Chew, 105 P
PITTOSPORAC

Benin iin Burk. ssp. ledermannii

Schod ees 10618

Ee horum Sinuata ae var. sinuvata, 11058

Paap sinuata Blume var. efuniculare Steen.,
10. 89

0655, 106

POLYGALACEAE
Polygala paniculata L., 10834

OLYGONACEAE
Muehlenbeckia platyclada (FE. Muell.) Meissn., 10934
estes chinense L., 10
PROTEAC
Helicia cf. Vee Fv.M., 11103
Helicta microphylla Diels, 10609
Helicta obtusata Sleumer, 10638

Helicia oreadum Diels, or aff., 10917

RHAMNACEAE
A JHhje : / Lr

ex Endl

,5R from

Camp 3
Ziziphus papuanus Laut., or Z. djamuensis Laut.,
‘rom Camp 4

Cee)
ROSACEAE
p dalgrbink

K. Schum. & Laut ) Kalkman,

11037
Prunus puller (Koehne) Kalkman, 10608
tla Kalkman, 10742
, 10588
ae pr aie Merrill & Perry, 10451, 10563
Rubus moluccanus L. var. moluccanus, 11013
a moluccanus L. var. obtusangulus Mig., 10453
Rubus trigonus Kalkman
RUBIACEAE
Atrosperma ramuense Laut. & K. Schum.,
Argostemma sp., 10

ae Se ies

10905
385; not A. sears
Dolicholobinm sa lobum K. Schum

Gardenia pallens Merrill & Perry, He oe
Hedyotis co aa 4 Br., 10965

Hedyotis pubescens Valeton, 11080
Hydnophytum radicans Becc., 10943

ee tum virgatunm ee. 11069

Ixora dolichothyrsa Brem., 10945

Nes umbellata L. var. ee Valeton, ne
Mus ssaenda oreadum Wernham, 10867; det

Mason is Wernham, 11097; conf.
ee: ideon

My Bl x Korth 278
iy Lyra ans malic Huxley & Jebb, ee 20
Myrmecodia schle

re ValesOnt 0720

ertera granadense <L. f.) Druce, 10504

Ophiorrhiza aff. 2amoena Valeton, 10991, 11019
Opbiorrbiza debruynii Valeton, 10382

eg Be lo Valeton, 109 20
ta platyc 1098.

‘nanifrutex group’, 10662

Psychotria olivacea Valeton, 11102

Psycho ie he sai dae & K. Schum.)
Valet

Bhai ee Sohmer, 10768
Psychotria valetoni lca: 10399, 10941

ellata ee or aff., 10830

Timonius beleiie Merrill & Perry, 10587

Timonius aft. xanthocarpus (oo & Perry, 10418

ta bernaysii Fv.

Un
U pe brittanicum Seas. or aff., 10756

RUTACEAE
Acronychia ledermannii Laut.,

10717, 10750
44

, 10606; small ie ate leaves,

Melicope sp. B, 106 : : eos petiolate leaflets
Melicope sp. C, 10823; gestalt like Melicope

780
mucronata, leaflets villous, sessile, to 37 cm
Meliopesp, D, 10892; glabrous obovate leaflets
SABIACEAE
Sabia pauciflora Blume, 11026
Meliosma eee (Roxb.) Maxim. ssp. bamilis

(Merri Perry) Beus., 10488
bie oles any ) Maxim. ssp. macrophylla
rrill) Beus., 11107
SANTALACEAE

Cladomyza cuneata Danser, 10436
SAPINDACEAE

Arytera aft. multijuga or lasting 10976
8

=)

Ps
a

‘©

Cupantopsis macropetala R
it 094A
Pometia pinnata Forst. & Forse. £., SR from Camps

Guioa comesperma Radlk.,

3 and 4
Sarcopteryx crispata Welzen, 11101
SAPOTACEAE
Planchonella (Un abanes ) HJ. Lan
presumably be
Krause

as ee montic ‘i

SOLANACEAE
Solanum anfractum Symon, 10530; det. Symon
Solanum rvostellatum Merrill & Perry, 1/0663; det.
Symon

SPHENOSTEMONACEAE
Sphenostemon papuanus (Laut.
10731

Steen. & Erdtman,

STERC IACEAE
pes ampla Baker f., 10963
Sterculia ee (L aut.)
Sterculia ck. schuma dd
SYMPLOC
Symplocos cor 7 joe (Lour.) S. Moore, 10566
10415

) Mildbr., 77085
.) Mildbr., 77081

Symplocos cf. cochinchinensis (Lour.) S. Moore,

~
nat
=
>
=
=)
o>
=.
=
=
>
x
=
a
S
=
na
oe
—
o
WN

ome Os + eachinchinensis (Lour.) S. Moore var.
Nooteb., 10748, 10843,

10961

cf. Symplocos; small leaves, 2uniflorous, 10507
THEACEAE

Eurya ct. leptantha Diels, 10394

Eurya sp., aia nee group’, 1065
, 10571
Ternstroemia ee Fv.M., 10532, 10624
‘Ternstroemia cherryi (EM. Bail.) Merrill, SR between

tamps 3 and 4

Eurya tigang K. Schum. & Laut., 1042

Stipa 18(3)

THY MELAEACEAE

Phaleria macrocarpa (Scheff.) Boerl., 10430
TRIMENIACEAE

Trimenia papnana Ridley, 10393, 10402
ULMACEAE

Parasponia vigida Merrill & eh 10567

10918

Trema cannabina Lour.,

TICACEAE

Boehmeria platyphylla D. Don, s. lat., 11016
Boehmeria sp., 10522 (m); aC ypbolopbus

C ace prema H. Winkler, 77078
Debregeasia sp.,

ae wate ee er, 1089

Elatostema aft. belense Perry, fone 10994
Elatostema blechnoides Ridley, 10450, 10478
aa 9

&
>

Elatostema macrophyllum B romps n.
Elatostema mongiense Laut ea
cai morobense Perry, /

se War ee 10885, 10896,

| 102 2?

Elatostema tridens Perry,
11096 (f)
10974
11044

0475

Elatostema weinlandii K. Schum.,

Laportea decumana (Roxb.) Wedd.,
Leucosyke i Migistad (Poir.) Wedd.,
Pilea effusa H. Winkler, 10512
Pilea pane H. Winkler, 106
Pilea stenonenra A. Winkler, oe 10410

*Pilea ue nov. oo) 10559, 10740
Pipturi res (Forse. f.) Wedd., 10446 (m),

10500 i: 10990 (f)

cee ‘uals H. a or aff., JO5/1
“hew, 10858 (f), LO984

(m)
Procris frutescens Blume, 10901
Procris sp., 1098
VERBENACEAE
Callicarpa longifolia HJ. Lam, 1097-
Geunsia pentandra (Roxb.) Merrill, 10821
Stachytarpheta ae (Rich.) Vahl, 7/028
VITACEA
Cayratia geniculata Blume, or att., 11027
Cc ci japonica (Thunb.) Gagn., SR from Camp

Cissus avistata Blume, 10948

Cissus discolor Blume, SR from Camp 3
Tetrastigma lauterbachianum Gilg, 11033

WINTERACEAE
Zygogynum oligocarpum (Schler.) Vink, 11084
Zyeogynum cf. sylvestre (A.C. Smith) Vink, 70850

FAMILY INDET., /0425, 10431

TAKEUCHI, Bismarck-Ramu expedition in Papua New Guinea 781
ACKNOWLEDGMENTS

The 1995 Bismarck-Ramu RAP Expedition was funded by the Global
Environment Facility and the United Nations Development Program. My
studies in Papuasian botany are supported by the Liz Claiborne and Art
Ortenberg Foundation, which I gratefully acknowledge.

Michael Hedemark and John Opo Akis were responsible for coordina-
tion of survey activities. The Lae National Herbarium served as a principal
logistical base and provided the facilities for processing and identification
of collections. Figures 1-3 were reproduced by permission from the De-
partment of Environment and Conservation (through Samuel Antiko, as-
sistant director, Biodiversity Assessment Branch). N.H.S. Howcroft illus-
trated the new species in figures 4-5. Colleague J. Pipoly of the Botanical
Research Institute of Texas prepared the Latin diagnoses.

Taxonomic specialists assisting with plant identifications included O. Gideon
(Costaceae and passim), N.H.S. Howcroft (Orchidaceae), J. Pipoly (Myrsinaceae),
P.F. Stevens (Clusiaceae, Ericaceae), and D.E. Symon (Solanaceae). Reviewer
P.F. Stevens made many incisive and helpful comments. Dendrologists K.
Demas and J. Wiakabu gave assistance with difficult taxa and also provided
corrections to the text.

REFERENCES

Airy SHAw, H.K. 1973. New or noteworthy species of Antidesma (Stilaginaceae): HI. Kew
Bull. 28:269-281.

Airy SHAW, H.K. 1979. New or noteworthy species of Antidesma (Stilaginaceae): V. Kew
Bull. 33:423-427.

Airy SHAW, H.K. 1980. The Euphorbiaceae of New Guinea. Kew Bull. Addit. Ser. 8.

Bartow, B.A. 1981. Loranthaceae. In: Henty, E.E. (ed.), Handbooks of the flora of Papua

ew Guinea 2:206—254

Bartow, B.A. 1997. Loranthaceae. Flora Malesiana ser. I, 13:209—401.

Bren_er, B.M. (ed.). 1993. Papua New Guinea conservation needs assessment, vol. 2. Cor-
porate Press, Inc., Landover, Md.

BrEHLER, B.M. (ed.) 1997 at a Rapid Assessment Program Working Papers. A
biodiversity assessment of southern New Ireland, Papua New Guinea. Conservation
International, Washington DC.

BROWNSEY, 987. A review of the fern genus Hypolepis (Dennstaedtiaceae) in the Malesian
and Paci regions. Blumea 32:227—276.

CraveEN, L. 1980. A new v Rbndadenn (Ericaceae) from New Guinea. Notes Roy. Bot. Gard.
Edinburgh 38:141—

Fropin, D.G. 1982. ie. In: Royen, P. van, The alpine flora of New Guinea 3:3040—

3083

Gruss, PJ. and P.F. Stevens. 1985. The forests of the Fatima Basin and Mt. Kerigomna,
Papua New Guinea: with a review of montane and subalpine rainforests in Papuasia.
Australian Nat. Univ./Anutech, Dept. Biogeogr. & Geomorph., Research School of Pa-
cific Studies, Publ. BG/5, Canberra.

782 Sipa 18(3)

HAMMERMASTER, E.T. and J.C. SAUNDERS. 1995. Forest resources and vegetation mapping
of Papua New Guinea. PNGRIS Publ. no. 4, Canberra, CSIRO and AIDAB

Harty, T.G. and L.M. Perry. 1973. A provisional key and enumeration of species of Syzygiam
(Myrtaceae) from Papuasia. J. Arnold Arbor. 54:160—227.

Hepemark, M.,S. Hamirron, and W. Takeucut. 1997. Report on the first Bismarck-Ramu
biological survey with sociological and logistical comments. PNG Dept. of Environ.
and Conserv., Port Moresby.

Hourtum, R.E. 1981. Thelypteridaceae. Flora Malesiana ser. H, 1 (5):331-599.

Jouns, R.J. 1993. Biodiversity and conservation of the native flora of Papua New Guinea.
In: B. Beehler, ed. Papua New Guinea conservation needs assessment report, vol. 2, Corporate
Press, Inc., Landover, MD. Pp. 15-75

KALKMAN, C. 1965. The Old World species of Prunus subg. Lawrocerasus including those
formerly referred to Pygewm. Blumea 13:1—115

KaLKMAN, C. 1993. Rosaceae. Flora Malesiana ser. I, 11 (2):227-351.

Kena, H. 1978. Labiatae. Flora Malesiana ser. I, 8 (3 301-394.

Ku LANG, J., O. Genta, and L. BALUN. 1997. A study on the biological diversity of the Hagahai
area in the ag Province, Papua New Guinea. Unpublished; informally distributed

as photocopie

MANSFELD, R. 1995. Die Melastomataceen von ae Bot. Jahrb. Syst. 60:105—

Martuew, B. 1995. Editorial. Bot. Mag.12(2):5

Maxwell, J.F. and J.B. VetbKAmp. 1990. . on the Astronieae (Melastomataceae) - I.
Astronidium, Beccarianthus. Blumea 35:115—164.

PHILipson, W.R. 1979. Araliaceae-I. Flora Malesiana ser. I, 9 (1): 1-105.

PHILIPSON, WR. 1986. Monimiaceae. Flora Malesiana ser. I, LO (2):255—326

Royen, P. van. 1964. Manual of the forest trees of New Guinea. Part 2 - bee. De-
partment of Forests, Division of Botan

RoyEN, P. vAN. 1982. The a flora of New Ss ea 3: taxonomic part Winteraceae
to Polygonaceae. J. C r, Vaduz. Pp. 1233-24

SMITH, R.M. 1978. Alpina sect. Pycnanthus at 1ts oe Guises species. Notes Roy. Botanic
Gard. Edinburgh 36:273-292.

Soumer, S.H. 1988. The nonclimbing species of the ee ig Cabaret: ae) in New
Guinea and the Bismarck Archipelago. Bishop Mus. Bull. t 1:1-339,

TakEUCHI, W. and J. KULANG. 1998. Vegetation part 2: cans survey. In: Mack, A., ed.
A biological assessment of the Lakekamu Basin, Papua New Guinea. Rapid Assessment
Peas rking Papers No. 9. Conservation International, Washington, DC. Pp. 36—
S910)

Senn 7 en Pipoty. 1998. New flowering plants from southern New Ireland, Papua
New Guinea. Sida 18:161-168.

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Kew B Addit. Ser. 8 —162

Wain, 1922. Die en ies Penuasens, Bot. Jahrb. Syst. 57:501—608

WomersLEY, J.S. (ed.) 1978. Handbooks of the flora of Papua New Guinea. Vol. I. Melbourne

University Press.

STUDIES IN NEW WORLD AMARANTHUS
(AMARANTHACEAE)

JAMES HENRICKSON

Department of Biology
California State University
Los Angeles, CA 90032, U.S.A.

ABSTRACT

Amaranthus crassipes is recognized as consisting of two varieties: var. crassipes from the
West Indies, coastal Mexico-Texas-Louisiana, and northern South America, and var. warnockii
of the Chihuahuan mee region of Texas and Mexico. Amaranthus scler Baoraes contains two
morphs that are not taxonomically recognized. Amaranthus < texensis, considered of hybrid
origin Pee i i sereasipes ane A. SLEDGE, is describe s new. Amaranthus ee is

id A new species, Amaranthus
tamaulipensis, is described from southmost Teas Gua eastern Mexico. It is related to
dubius and the poorly known A. brandegei of Sinaloa. An enigmatic ee: x fucsonensts
is described from the Tucson Mountains of southern Arizona, with related populations occurring
in Sonora, Mexico.

RESUMEN

e reconoce a Amaranthus crassipes como constituida por dos variedades: la var. crassipes,
con distribucién desde la costa de México, Texas y Louisiana, a través del Caribe hasta e
norte de Sudamérica; y la var. warnockii, de la region del Desierto Chihuahuense. Amaranthus
scleropordes consiste de dos fases morfolégicas sin designacién de rango taxonémico ;
La notoespecie nueva Amar. a x texensis eae se origino de la ces en-
tre A. crassipes y A. scleropoides. No se consideraa A. berlandieri como diferente de A. ae
especie de mas amplia distribucién. Se describe as tamaulipensis, €
del este de México y la parte mds meridional de Texas. Esta especie esta ee con ay
dubius y con A. ee taxon sinaloense pobremente conocido. Se describe la enigmatica

Amaranthus X tucsonensis com I nueva de la Sierra de Tucson, del sur de Arizona.

Key worps: Amaranthus, Amaranthaceae, plant taxonomy, plant s atics, hybridization

Studies in the Amaranthaceae in connection with the Chihuahuan Desert
Flora have resulted in this paper on the systematics of the Amaranthus crassipes-
A. scleropoides and A. polygonoides groups. Two additional taxa are described
as new.

The Amaranthus crassipes-A. scleropoides group has dichasial clusters of flowers
borne at all nodes from the base of the plant to the tip. The inflorescences
are modified compound dichasia in which the terminal flowers are overtopped
by opposite, sinuous peduncles that bear one to a few pairs of persistent
bracts with mostly male flowers and terminate in pistillate flowers that are
again overtopped by similar paired, sinuous peduncles. As the peduncles

Sipa 18(3): 783-807. 1999

784 Sipa 18(3)
mature they become greatly thickened and indurated in some species and
trap the fruit between the expanded peduncles (Fig. 3A, D). Gray (1862)
considered thickened peduncles to be an abnormal character in A crassipes,
but Holzinger (1892) showed they were consistently produced in that spe-
cies, caused by a proliferation of expanded parenchyma cells that develop
thickened walls. At maturity, the entire inflorescence falls from the plant
and serves as a dispersal unit capable of flotation.

VARIATION WITHIN AMARANTHUS CRASSIPES

Amaranthus crassipes Schltdl. 1s a widespread New World species occur-
ring from the West Indies, Bahamas, Florida Keys, Arizona, Texas, Mexico,
Colombia, and Venezuela (Fig. 1)—its type locality is on St. Thomas in the
Virgin Islands. The species consists of decumbent-procumbent, glabrous,
monoecious annuals with ovate-obovate to lanceolate, long-petiolate leaves.
The flowers are produced at each node usually from the base of the plant
upwards in short dichasia 5-10 mm long (Fig. 2C, E). The dichasia de-
velop the aforementioned greatly thickened, indurate, strongly sinuous lateral
axes (Fig. 3A). The female flowers have 5 erect to spreading, narrowly spatulate
sepals chat are thickened and joined at the base (Fig. 3B). They are usually
shorter than the fruit. The fruit wall is closely adherent to the seed and indehiscent,
although it may develop a colored encircling line below the middle. It is
usually smooth below and somewhat tuberculate above and along the mar-
gins. The 2(—3) styles are joined together into a distinctive lyrate-shaped
neck before they taper and diverge as slender tips (Fig. 2A—C). The male
flowers are produced on the sides of the developing inflorescence axes. They
mature and fall away before the inflorescence axes thicken and they have 5
membranous sepals and three anthers.

Throughout the Caribbean region the plants are initially erect but usu-
ally become procumbent weeds in disturbed places. They usually have ovate,
broadly ovate to obovate leaves that may either be uniformly small or, in
fast growing plants, large in size. (Fig 2C). Similar ovate-leaved plants also
occur in northern South America, the Yucatan Peninsula, northeast Mexico,
coastal Texas, Louisiana, Key West (Florida) as well as Sonora and Arizona.
In contrast, plants in inland Texas and in the Chihuahuan Desert region,
develop narrower, more oblanceolate, grayish leaves (Fig. 2E). They also tend
to be more erect or decumbent in growth habit, features that are shared
with the related A. sc/eropoides. The grayish color of the leaves also make the
reticulate pattern of the Kranz structure less conspicuous. In 1944 I.M. Johnston
described these inland plants as Amaranthus warnockii, distinguishing the
species by its elongate, somewhat thinner oblanceolate leaves, its less elon-
gate, more slender stems, and smaller more compact cymes conspicuously
crowded at the stem bases. Except for the narrower, more oblanceolate-lan-

poet

—

HENRICKSON, Studies in New World Amaranthus 785

Amaranthus crassipes
@ var. crassipes
© var. warnocki

| SCALE

eee
CONIC PROJECTION

— —, -—————— 1, —-—-- =

Fic. 1. Distibution of Amaranthus crassipes var. crassipes and var. warnockii.

ceolate leaves, all the other features noted by Johnston also occur in occa-
sional plants of A. crassipes throughout its range. In all technical characters
of flowers, fruit and inflorescence, A. warnockii and A. crassipes are identical,

Recognition of these two taxa at the species level is considered unjustified
as the distinguishing features are minor and to some extent inconsistent
and recognition would create poorly differentiated microspecies within this
highly variable genus. But overall the group of specimens referable to A.
warnockii can be recognized solely on the basis of the narrower, more oblan-
ceolate, grayish leaves. Because of this A. warnockii is here retained at the
varietal level as it represents a geographically based subunit of the species
(Fig. 1). Correll and Johnston (1970), in contrast, considered A. warnockii
synonymous with A. crasszpes. The collections from southern Arizona and
adjacent Sonora, however, have distinctly ovate leaves and fall within var.
crassipes. A formal description of A. crassipes follows.

AMARANTHUS CRASSIPES

Low spreading, taprooted, procumbent to decumbent, basally branched,
monoecious annuals, sometimes with the central stems erect-ascending; stems
glabrous, striate, stramineous, moderately branched, 1—4(—9) dm long; in-
ternodes 1.5—4 cm long below, reduced to S—15 mm long distally. Leaves
alternate, ascending, (12—)30—55(—78) mm long, the petioles slender, (S—)
10—25(-47) mm long; leaf blades broadly to narrowly obovate to ovate, to

786 Sipa 18(4)

oblanceolate or elliptical, (7—)20—30(-50) mm long, (S—)10—20(—28) mm
wide, obtuse to rounded, usually retuse with the midvein short-excurrent
at the tip, narrowly acuminate-cuneate at the base, the margins entire to
erose-crisped, whitish, the veins curved towards the tip, but not extending
to the margins, distinctly whitened beneath, the lamina glabrous, with distinct
Kranz internal anatomy. Flowers borne in axillary, congested, tan or rarely
dark brown, glomerate dichasia 5-9 mm long usually borne at all nodes
from the base of the plant upwards, the dichasia decussately branching below
each pistillate flower, the branches strongly sinuate-curved, becoming thickened
and indurated, 0.9-1.2(-1.6) mm thick and crowding-enclosing the de-
veloping fruit; bracts and bracteoles broadly deltate, 0.5—O.9 mm long, green
along the midrib, the margins scarious, broadly attached at the base, per-
sisting on the sides of the thickened dichasia. Staminate flowers produced
on the sides of the developing dichasia, with 5, subequal, lanceolate sepals
1.2—1.6 mm long, these membranous except where green along the mid-
rib, acute; stamens 3; anthers 0.7—0.9 mm long; filaments 1.5—2 mm long.
Pistillate flowers born at each branch of the dichasium; fruiting sepals 5,
with 2 at each margin of the fruit and with a smaller sepal on one ee 2on
the other face, the sepals distinctly clawed or merely spatulate, 1.4—2.3 mm
long, the claws narrowly linear, 0.1—0.2 mm wide, keeled, are to
ovate, acute, slightly keeled, conduplicate, sometimes slightly reflexed tips
0.5—l mm long and nearly as wide, the tips each with a green in a trullate
patch medially and with membranous margins, the sepals expanding to become
spongy-indurate, tan, united and truncated around the fruit base. Fruiting
ovary compressed orbicular-obovate, 1.3—1.5 mm wide and long (exclud-
ing the style), the fruit wall somewhat thickened, closely surrounding the
seed, indehiscent or in some plants with a darkened line in the lower half,
usually but not consistently smooth below, distinctly tuberculate above and
along the margins, topped with the persistent style; che styles 2(—3), about
1.5 mm long, connate and erect for 0.2—0.5 mm below, the yellowish lobes
erect or usually lyrate, curving outward, then inward as they taper, termi-
nating as slender outwardly curved appendages, the papillate-stigmas ex-
tending all along the inner surface of the styles onto the terminal slender
portions. Seeds compressed-obovoid, 1.1—1.3 mm long, 0.9-1.1 mm wide,
0.6-0.7 mm thick, shinny, dark brown-black, the margin obtuse to some-
what rounded, the surface shiny, with a visible cellular surface pattern. With
two varieties:
Amaranthus crassipes wane var. crassipes (Figs. 2A—C, 2A—B). Linnaea
3:757. 1831 (as Amarantus). § 11.) Moq. in DC. Prod. 137:271.
1849; Enxolus crassipes (Schltdl. \ Fidion:. Bol. Acad. Nac. Cérdoba 4:13. 1881. Type:
WEST INDIES. St. Thomas Island. (Ad rivulos insulae St. Thomae, 1826-1828)

HENRICKSON, Studies in New World Amaranthus 787

E

Fic. 2. Fruits and foliage of Amaranthus crassipes. A-C. A. crassipes var. crassipes. A, Mature

a note characteristic lyrate styles and clawed sepals that are united at the base ee

Co., Tex., Rayon 658, TEX). B. Mature fruic eee: Box 573, NY). C. Characteristic

ovate ee bi: ee hicola, Fla. » Chapman Herb. 3 383, (NY). ). D-E. A. crassipes var. warnocktl.

D. Mature fruit (Pre Co. Tex., Hinckley 129, SRSC). E. Characteristic oblanceolate leaf

4 = ee Co. iy ee & Johnston 17209, SRSC). White bar in A = 1 mm for A,
1 for C. E.

is J.m. (HOLOTYPE: HAL). Dr. U. Braun has sent a xerox copy of the holotype
HAL, deposited TEX
Scler opus ies Schrad., Index sem. Hort. gott. 1835; (reprinted in: Linnaea 11:
-ber. 89. 1837). Tyre: unknown.

Plants nae erect when young but becoming procumbent, occasion-
ally decumbent, /eaf blades ovate to broadly ovate, sometimes nearly orbicular or
obovate, (7—)15—30(—50) mm long, (S—)10-15(-28) mm wide, the ower surface
green to yellow-green, distinctly marked by the whitish, arcuate veins.

Weed in roadsides, waste grounds, fields, pastures, in clay, silty, some-
times sandy flats, beaches, and rocky slopes in shaded or open habitats, widespread
in the Caribbean region, south Florida, coastal Louisiana, Alabama (adven-
tive), Texas, northeastern Mexico south to northern South America (Columbia,
Peru), from sea level to 250(—1250) m (Fig. 1). It also occurs in Arizona,
Sonora, Chihuahua at higher elevations. Collected throughout the year but

788 Sipa 18(3)

expected to be vegetatively active during hot months due to C4 photosyn-
thetic pathway.

Amaranthus crassipes Schltdl. var. warnockii (I.M. Johnston) Henrickson,
comb. nov. aa ig. 2D-E). Basionym: Amaranthus warnockii 1.M. Johnston, J. Arnold
Arbor. 25:153. 1944. Type: MEXICO. Coanuia: a mi SE of Ocampo, low place
near mogote on plain: 8 Sep 1941, 1M. Johnston 8886 (HOLOTYPE: GH!).

Plants initially erect when young, the lateral branches becoming decumbent,
ascending terminally; leaf blades oblanceolate to narrowly oblanceolate, (12—)16—
33(—44) mm long, (4—)6—10(-16) mm wide, the swrfaces more glaucous-green,
the veins of the lower surface slender, not conspicuous.

Open silty flats, mesquite thickets, muddy areas to 1300 m elevation, in
south-west Texas and in the Chihuahuan Desert in Coahuila and eastern

ee

—

vihuahua (Fig. 1).
AMARANTHUS SCLEROPOIDES

In 1895 Uline and Bray described a second species related to A. crassipes
differing in its circumscissile, smooth utricle and 3 style branches. Uline

and Bray considered Amaranthus scleropoides intermediate between A. crassipes
and A. graecizans auth. non L. (now A. albus L.), the latter also having 3 styles
and circumscissile utricles. Vegetatively and in inflorescence structure, A.
scleropoides is very similar to A. crassipes, but A. scleropordes is a taller, bushier
plant. While its mature utricles are uniformly circumscissile, the pistillate
flowers are somewhat variable in sepal structure, style number and other
features. Mature sepals vary from 1.2—2.5 mm in total length. As in A. crasszpes,
two sepals are usually distributed at each edge of the mature fruit, with a
two on one face and a one smaller sepal on the other face. The sepals are
clawed and vary greatly in development. In some specimens the blade por-
tion is erect, green, rounded, apiculate and rather flattened, however, in others,
including the type collection, the tips are distinctly thickened, keeled, con-
duplicate, recurved, and sharp pointed, with a central green rhomboid patch
(Fig. 4C). Styles vary from two to three (Fig. 4A—B), or rarely five (1-flower
on Cory 2933 GH), with some plants being consistently two styled and others
with varying numbers. In flowers with two styles, the styles are located directly
over the margins of the compressed ovary and as the fruit mature the styles
develop distinct decurrent ridges that extend down the fruit margins. When
styles are three, the third, often reduced style, develops on one of the faces,
again with a distinct decurrent base. The surfaces of the utricle may be smooth
or tightly turberculed.

Among collections observed, two distinct forms are recognizable based
on differences in the development of the styles and upper ovary wall. The
most common form has distinctly spongy-thickened style bases and the upper

HENRICKSON, Studies in New World Amaranthus 789

Fic. 3. Amaranthus crassipes and A. scleropoides. A-B. A. crassipes var. crassipes. A. Mature
dichasial inflorescences have spongy-thickened, indurated lateral branches; the external bracts
subtended male flowers, the female flowers are located at the base and tips of the inflores-
cence branches (Jamaica, Proctor 23937, NY

). B. Mature fruit-calyx Jamaica, Proctor 23937,

’). C-E. A. scleropoides. C. Young plant with characteristic narrow leaf blades, axillary
inflorescences. D. Mature inflorescence. E. Mature fruit and calyx (all Williams Co., Tex.,
Walcott 303, TEX). Magnifications in A & B hold for D & E.

790 Sipa 18)

utricle is indurated and stramineous in color (Fig. 4A—B). The styles are
papillate (stigmatic) along their inner margins and the papillae continue
onto the slender terminal lobes. A second form does not develop expanded,
indurated thickenings on the style bases and upper fruit and the two or
three styles are thus clustered at the top of the fruit (Fig. 4C). The two forms
both occur throughout the range of the species—they do not sort out geo-
graphically (Fig. 5). Because of this, and because this variation does not
correlate with any other morphological or ecological variation, they are here

recognized as unnamed forms. A description of the species follows.
Amaranthus scleropoides Uline & ee Bot. Gaz 19:316. 1894. hee
°—-E, 4A—-C, 5). Type: U.S.A. TEXAS: “Texas, Western Texas to El Paso,” Wright
582: (LECTOTYPE: GH; ISOLECTOTYPE an Notation on lectotype by I.M. Johnston:
“Texas: Val Verde Comune), Wright feld number 798, Amaranth, Devil’s River, sum-
mit of hills, 2 Aug 1849.” The type has sepals with a sharply divergent

tips; the style base is not inflated as in some collections

Very similar to Amaranthus crassipes vegetatively and in inflorescence and
flower organization, but typically more erect, bushy, 2—5 dm tall and often as
wide, with the lower stems ascending or decumbent and ascending to erect
distally. Leaves with slender petioles (4—)10—25(—47) mm long; leaf blades
lanceolate, oblanceolate to narrowly trullate, (S—)15—25(—42) mm long, (2)
5—10(—12) mm wide, glabrous, somewhat glaucous. Flowers borne in thickened,
sinuate dichasia produced from the base to the top of the plant; staminate
flowers with 5 sepals, 3 stamens. Pistillate fruiting sepals 5, with 2 at each
margin of the fruit, two sepals on the one face, with one smallest sepal on

—

the other face, the sepals moderately clawed to narrowly spatulate, in fruit
(1.2—)1.5—2.2(—2.5) mm long, the claws 0.2—0.7 mm wide, the distal sepal
blade typically broader, erect or variously reflexed, + keeled and conduphi-
cate, + ovate-lanceolate, acute, with a small or large, central, green narrow
to trullate green patch to 0.5—1 mm long, the margins membranous, the
sepals expanding and becoming spongy-indurate, tan, and united around
the fruit base. Fruiting ovary compressed orbicular-obovate, 1.1—1.3 mm
wide and long (excluding the styles), the wall circumscissilly dehiscent, with
a distinct brownish medial line, the body smooth or becoming closely turberculate
above and less strongly tuberculate below the line of dehiscence, the styles
2—3(—5) [the third (fifth) one(s) smaller when present}, arching outward from
the upper margins of the fruit wall, the style bases continuing as decurrent
ridges down the fruit wall, the lower portion of the styles and adjacent dis-
tal fruit wall (but not extending down to the line of dehiscence) often spongy-
thickened, inflated, or the distal fruit wall and adjacent style bases only slightly
inflated and ascending; the stigmas papillate, extending along the inner margin
of the style lobes and continuing as slender tips for 0.4—0.5 mm. Seeds compressed
mm long, 0.85—0.96 mm wide, 0.6—0.7 mm thick, dark

—
on

obovoid, |.0—1.

HENRICKSON, Studies in New World Amaranthus 79)

brown—black, the margin obtuse to somewhat rounded, the surface shinny,
with a slightly visible cellular surface pattern.

A NEW TAXON FROM SOUTHERN TEXAS

Within the complex of A. crassipes and A. scleropoides, a series of speci-
mens from southern Texas are characterized by an erect-ascending growth
habit and what appear to be circumscissile utricles as in A. scleropoides. However,
the utricles are thick walled, weakly and closely tuberculate and are actu-
ally indehiscent as in A. crasszpes although a distinct medial line is present.
As in A. crassipes, the styles are united at the base into a distinct neck before
they separate, but the 2—3 styles have distinct decurrent ridges that extend
down the utricle wall as in A. scleropoides (Fig. 4E). These specimens also
have rather narrowly lanceolate to linear-lanceolate, long petioled leaves,
and they tend to have longer, more strongly reflexed pistillate sepals of a
type sometimes found in A. sc/eropoides. The specimens, some of which are
sterile and produce no mature seed, appear to represent hybrids or deriva-
tives of hybridization, perhaps between A. crassipes and A. scleropoides. These
collections are all from loose red sands to sandy loams in southern Texas in
Duval, Webb, Hidalgo, and Atascosa counties of southern Texas, with one
collection from adjacent Tamaulipas. They are of sufficient distinction to
be recognized as a separate nothospecies.

tay a x texensis Henrickson, nothosp. nov. ees 4D, 5). Tyee:
S.A. TEXAS. WEB BB Co.: in red sand at roadside on Hwy. 83, 13 mi NW of Webb,
ou 1957, D.S. Correll & 1M. ee 18101 (HoLoTYPE: LL!; isorype: GH!).

nN A +h ] ed leh _ py GP 3 Faye eo! —ee erees ro

semilis sed differt Basibne stylaribus collum distinctum 0.2—0.4 mm longum formantibus
(ut A. crassipe) et sepalis florum pistillatorum valde reflexis ad apices.

Annual herbs 10—35 cm tall; stems erect-ascending to decumbent, with
scattered obscurely gland-tipped hairs to 0.3 mm long, glabrate. Leaf blades
linear-oblanceolate, (S—)13—25(—40) mm long, 2—6(—8) mm wide, obtuse-
ee with the midvein excurrent at the tip, narrowly cuneate, the
mar xtending well down the (3—)5—15(—20) mm long petiole, the margins
which eon the blades green to gray-green with the veins impressed
above, stramineous and raised beneath. Flowers borne in short, thickened,
axillary dichasial cymes produced at each node of the plant from base to
tip, the cymes 3.5—6 mm long, the cyme axes sinuate, stramineous, spongy-
thickened, indurate, 0.5—1 mm thick, bracteate, the bracts |—-1.5 mm long,
green medially in a diamond pattern with broad clasping scarious bases,
the staminate flowers early deciduous, the dichasial cymes bifurcating at
the bases of each pistillate flower and terminating in pistillate howers; staminate
flowers: sepals 5, narrowly ovate, 1.5—1.8 mm !ong, membranous except

along the excurrent midvein; stamens 3; mature pistillate flowers: sepals 5,

Sipa 183

792

Amaranthus feseroporaes. A-B. Fruit

Fic. 4. Photographs of mature fruit and calyces. A-C. .
Cameron Co., Tex., Ramyon 2397, F). C. Fruic with
York 46249, F). D. A.

texensis. Mature fruit; note oe base, sharply recurved sepals (Webb Co., Tex.,

with inflated s tyle bases, 2 or 3 styles

non-inflated style base, note recurved sepals (McLennon Co., Tex.,

& Johnston 1S1O1, LL-type A. polygonoides. Mature fruit-sepals, note sepals have 3 veins
and are inflated and indur aa at base (Santo Sate. Ekman | es , US). EA. brandeges.
, México, Brandegee

Mature calyx with reflexed, broad sepals, inflated top of
y.0.-Typr, UC). € - — Note spachulate, greet Gena inflated Seen vameron
, Runyon 37 a tucsonensis. Mature calyx and pericarp top (Pima , Ari

Devender 88- 8 c ARIZ). White bar in A = | mm holds for all figs.

HENRICKSON, Studies in New World Amaranthus 193

|
Amaranthus scleropoides

Fic. 5. Distribution of Amaranthus scleropoides, A. tamaulipensis, A. X texensis, A. X tucsonensis
in southwestern U.S.A. and México.

clawed, 1.5—2.1 mm long, to 0.3—0.6 mm wide below the acute tips, spongy,
thickened and all conjoined at the base, the distal portion expanded, strongly
spreading, becoming nearly horizontal below the spinose tip, the base and
claws stramineous, sometimes ciliate with slender gland-tipped hairs, the
expanded tips with an elongate green patch 0.9-1.3 mm long, with nar-
row scarious margins. Fruiting ovary vertical, the wall initially smooth and
with a medial encircling line, but in age becoming thickened, bullate-tu-
berculate in the upper half, remaining smooth below but indehiscent, the

794 Stipa 18(4)

styles 3(—2), joined into a vertical neck for 0.2—0.5 mm before the tapered
styles gradually arch outward, when styles 3, with two styles larger than
the third, the styles papillate along their the inner surfaces.

The new nothospecies occurs with A. crasszpes in Webb Co., Texas and
some collections (i.e., McCart 8559) are mixed with that species. ee ee
x tenensis is similar to A. scleropoides in most features, but stands apart in
the distinctive narrow neck produced by the combined style bases (Fig. 4D)
and by the thickened utricle wall-in these characteristics it relates to A.
crassipes. The strongly reflexed sepals, are overall of similar structure to those
found in some specimens of A. sc/eropoides, but are much thicker, as in A.
crassipes. The pistillate sepals of A. crass/pes are erect and may or may not be
reflexed at the tip.

Additional collections: U.S.A. TEXAS. Atascosa Co.: State Hwy. 140, 4 mi W of Charlotte,
3 Nov 1962, McCart et al. 8538 (TEX). Duval Co.: Texas Hwy. 359, 6.5 mi E of Bruni, 4
Nov 1962, McCart et al. 8559 (TEX, LL—mixed with A. crassipes var. crassipes). Hidalgo
Co.: McAllen Ranch, 26 Jul 1951, Tharp, Follansbee & Thompson 51-1628 (TEX). Webb
Co.: State Hwy. 359, | mi E of Acris. 4 Nov 1962, McCart et al. 8595 (LL); Farm Road
1472, 10 mi NW of Laredo, 23 Nov 1962, So/is 93 (TEX). MEXICO. Tamaulipas: 34 mi
S of ieeaimeiens on road to San Fernando, 14 Sep 1960, Johnston & Crutchfield 5488C (TEX).

AMARANTHUS POLYGONOIDES VS. AMARANTHUS BERLANDIERI

Amaranthus polygonoides L. and A. berlandieri (Moq.) Uline & Bray are similar
species readily distinguished from other Amaranthus by a number of fea-
tures (Fig. 6). In both the flowers are borne in tight axillary dichasial clus-
ters at each node that become distinctly crowded at the uppermost nodes.
The peduncles are not thickened at maturity. The pistillate sepals are thin
and distinctly three veined in the claw area, with the central vein extend-
ing into the expanded, obtuse-acute, spreading blade area. The sepals are
united at the base and this basal area expands via cell enlargement to form
a spongy, somewhat indurated flotation device. The fruit wall is thin, cy-
lindrical, inflated, dehiscent or not, and is topped by three expanded style
bases that are exserted above the spreading sepals.

Uline and Bray (1895), Standley (1917), Correll and Johnston (1970)
and others have recognized both A. polygonordes and A. berlandieri as dis-
tinct species, basically repeating the characteristics given Uline and Bray
(1895) and modified by Standley (1917) in his key as noted below:

_

Utricle circumscissile; leaf blades suborbicular to ovate or oval, the leaves not

crowded; West Indies, Florida, Texas, Mexico, northern South America.
A. polygonoides
Utricle indehiscent; leaf blades oblong- lanceolate, the leaves crowded at the

ends of the branches; central-west Texas. . berlandiert

Moquin-Tandon (1849), in his treatment of the “Amarantaceae” (sic) in
DeCandolle’s Prodromus, recognized the above two species in distinct gen-

HENRICKSON, Studies in New World Amaranthus 795

Fic. 6. Variation in A. polygonoides. A-C. Typical Caribbean aan showing obovate leaf
blades. B. Mature fruit with subtending calyx, note 3- Be sals. C. Mature inflated,
dehiscent fruic (all Monterrey, México, eee M375, TEX). B - Typical Texas cise
showing more lanceolate leaf blades. E. Mature fruit wah subtending calyx, note 3-veined
sepals. F Mature inflated, non-circumscissile fruit. (all Cameron Co., Tex. Ce
17935, LL). Magnifications as indicated.

era in separate subtribes of his tribe Achyrantheae. He recognized taxon
polygonoides in subtribe Aerveae, (with indehiscent utricles) as Amblogyna
polygonoides Raf. (monoecious, staminate flowers with 3 sepals, 3 stamens,
the pistillate calyx being subglobose-funnelform, 5-parted, and the inflorescence
axis not becoming thickened as in A. crassipes). Moquin-Tandon’s newly described
taxon berlandieri was placed in subtribe Amaranteae (with utricles circum-
scissile) as a new genus Sarratia next to Amarantus (sic), differing in having
staminate Howers with 5 sepals and pistillate calyces funnelform-urceolate.
In his description he noted Sarratia berlandieri had 5 anthers per male flower
and circumscissile utricles.

796 SipA 18(4)

Moquin-Tandon made several errors in characterizing these taxa. In both
taxa, staminate flowers have 5 (rarely 4) sepals and both taxa have only 2
anthers per staminate flower. My observations showed that, contrary to his
data, the fruit of specimens referable to A. ber/andieri are indehiscent (not
dehiscent as he stated) while those of taxon pol/ygonoides are usually, but not
consistently dehiscent (not indehiscent as noted by Moquin-Tandon). These
errors were caught by Gray (1861, p. 168) who, in expanding the genus
Amblogyne to include two species previously placed in Sarratia, included taxon
berlandier: within his Amblogyne polygonoides noting that the specimens upon
which taxon berlandieri was founded were very poor. In contrast, both Uline
and Bray (1895) and Standley (1917) recognized both taxa within the ge-
nus Amaranthus. Thellung (1914), in contrast, treated berlandieri as a sub-
species of A. polygonoides.

During an initial study of these taxa it was noted that the characters in
Standley’s 1917 key did not work consistently and several specimens showed
characteristics of both taxa. Regarding fruit dehiscence, an analysis of 118
collections from F, GH, NY, TEX-LL, and US revealed the following. Specimens
attributable to Amaranthus berlandieri are mostly indehiscent (Fig. 6F), but
occasional specimens (2 of the 42 collections examined) did show a distinct
encircling line below the rugate cap of the utricle. In the 76 collections of
A. polygonoides with mature fruit examined, 36 were noted to be actually
circumscissile (Fig. 6D) or at least had developed a colored line encircling
the fruit wall below the tuberculate cap (however, the fruits may or may
not dehisce along this line) and 35 (49 percent) were clearly indehiscent.
Indehiscent utricles occur in collections from coastal Texas, Cuba, and throughout
the Caribbean region, often in localities where other collections clearly have
circumscissile fruits. In all instances care was taken to insure that the fruit
walls were mature when the dehiscence character was scored.

As noted in Standley’s key, there are some recognizable differences in leaf
shape. Most specimens from the Caribbean region to coastal Texas have ovate
to broadly ovate, rarely obovate (Fig. 6A), slightly more greenish leaf blades
and many specimens from interior Texas and Mexico have more narrowly
trullate or trullate-lanceolate (Fig. 6D), somewhat bluish-gray leaf blades.
To illustrate this a series of leaf-blade length-width ratios were taken. The
ratios of specimens from the Caribbean and coastal Texas attributable to A.
polygonoides range from 1.3 to 2.6 times longer than wide. Those of inland
areas attributable to A. ber/andieri have leaf-blade length-width ratios ranging

rom 2.1 to 4.2 times longer than wide. The problem is that when plants
are mapped out based on these leaf characteristics we find several collec-
tions from Sutton, Edwards, Tom Greene, Terrell, Presidio and Pecos counties
in Texas, as well as collections from Coahuila, that clearly have broad leaves

HENRICKSON, Studies in New World Amaranthus 797

as in taxon polygonoides. In many cases nearby specimens have narrow leaves
as in taxon berlandier?. Recognition of two taxa on the basis of this charac-
teristic would be quite arbitrary. Likewise leaves and inflorescences are crowded
at the distal stems in both taxa and the taxa could not be separated on the
basis of this characteristic. In fast growing stems, the leaves in both taxa
are well separated by long internodes.

As the distinguishing characteristics of these taxa are very weak and variable,
the two taxa are combined into a single variable species with recognition
that there are some trends present, but they are insufficient for nomencla-
tural recognition. Thus only one species is recognized from this complex as

ollows.

Amaranthus ea nase ee 4E, 6, 7), Pl. jamiac. pug. 2:27. 1759.
Roemeria polygonoides (L.) M h. Meth. 314. 1794; eres ee (LE) Rat..
Fl. Te al ee 1837; Aiea Sey th, FI. 2:144 oe
Sarr Mogq 13(2):270. 1849, as synonym. ee PE: JAM
Linnaeus’ 1759 proto referenced ‘ ce jatny do te Oe te. EL eae .
Madera Jamawa 1:144 tab. 92, fig. 2. ee wets the taxon was designated by a
polynomial “Blitum Vos virde, 5 ex veridt & eatum, polyanthos”. Linnaeus
only saw Sloane’s figure and not thee specimen from which it was drawn, hence the
specimen (Herb. Sloane 2:116, BM), can not be designated as lectotype. I her
designate the illustration in Voy, Madera Jamaica t. 92, fig. = as lectotype. The ee
Sloane plate 92, and the Sloane specimen, from which it was drawn, are at BM (C

Jarvis, pers. comm.). Sloane (L.c.) notes: “It grows in hard Chiy grounds, and amongst
Rubbish, every where about the Town of St. Fago de la Vega.” St. Fago de la Vega is
now known as Spanish Town. Elizabeth A. Kellogg, F/. Lesser Ant. 4:160. 1988, treated
the Sloane specimen as “Type,” however, for reasons noted above, it may not serve as

lectotype
Amaranthus verticillatus Pavon, Mog. in DC. Prod. 13(2):270. 1849, as synonym
Sarratia berlandieri Mog. in DC., Prod. 13(2):268. 1849; Amaranthus berlandieri Gisa: )

Uline & Bray, Bot. Gaz. ee 1894. Amaranthus polygonoides subei berlandieri (Moq.)
Thell. in Ascherson & Graebener, Syn. es FI. 5(1):352. 1914. Type: MEXICO.
Inter S. Fernando et Matamoros, eee 2279 (HOLOTYPE: G-DC not seen; IsoTYPE: GH!).

Low spreading to erect-ascending, monoecious annuals 2—4 dm tall; stems
branched below, the lateral (or all) branches decumbent, sometimes ascending,
1-4 dm long; internodes 2—5 cm long below, reduced to 2—5 mm long distally,
stramineous, striate, puberulent-villous with crisped moniliform hairs 0.2—
0.5 mm long, the lower stems glabrate. Leaves alternate, ascending, often
crowded, larger in the upper half of the plant; petioles (2—)5—20(—40) mm
long; leaf blades ovate to trullate-lanceolate, sometimes obovate or lanceolate,
(4—-)8—25(—42) mm long, (2.4—)5—15(—23) mm wide, rounded, obtuse or
emarginate with the midvein excurrent at the tips, narrowly to broadly cuneate
at the base, glabrous, the margins entire to erose, the veins and margins
whitish beneath, the lamina with distinct Kranz internal anatomy, green

798 SipA 18(3)

Fic. 7. Distribution of Amaranthus polygonordes,
&

to grayish in color. Flowers borne in axillary congested globose dichasia 3—
8 mm in diameter, these remote on the lower branches but becoming ag-
gregated as the internodes shorten above, the inflorescence axes thin, to 2
mm long; bracts and paired bracteoles lanceolate, 1—-1.5 mm long, mem-
branous, curved and cupped below, attenuate often with a long excurrent
midvein at the tip; staminate flowers: sepals (4—)5, + unequal, 0.7—1.3 mm
long, membranous except for the midvein; stamens 2, anthers 0.4—0.6 mm
long, filaments moderately thickened, to 1 mm long; pistillate flowers: mature
calyx (1.6—)2.3—3.0 mm long, the sepals united and becoming spongy-thickened
in the lower third forming an expanded float 0.9—1.2 mm long and wide
that slightly extends over the subtending broad-tipped pedicel, the sepa-
rate lobes clawed, spathulate, the claws 0.3—0.4 mm wide, notably 3-nerved,
with scattered slender gland-tipped hairs, the lobes to 0.6-1.2 mm wide,
erect to recurved-spreading, rounded to acute at the tips with the midvein
sometimes excurrent, membranous except for the 3 veins, the lateral veins
straight, slightly spreading outward, diminishing below the margin, the
midvein strongest, unbranched, when young overlain with a narrowly oblan-
ceolate central green patch. Mature ovary wall (the utricle) inflated, adnate
to the spongy calyx in the lower third, the mid portion smooth, compressed,
0.70.9 mm long and wide, sometimes developing a darker encircling line
just below the expanded tip (where dehiscence may occur), the distal tip

HENRICKSON, Studies in New World Amaranthus 799

portion variously roughened, sometimes distinctly expanded at the base,
capped with the narrower, fused, obconate, expanded style bases 0.4—0.6
mm long and wide, the styles 3(—2), the papillate slender terminal portions
0.5 mm long. Seed vertical, lenticular, 0.8—0.9 mm long, + 0.7 mm wide,
0.35—-0.5 mm thick, reddish-brown, smooth, shiny, the margins thin.

The species ranges throughout the Caribbean region to coastal Central
America, Key West Florida to central and western Texas and adjacent Mexico
(Fig. 7) occurring mostly in disturbed habitats. Also cultivated in Germany,
Italy and Egypt (Thellung 1914). Collections from Sonora and Sinaloa with
more erect, slender habits and similar staminate and pistillate flowers but
with the central vein of the pistillate sepals conspicuously branching are
referable to Amaranthus venulosus 8. Wats.

A NEW AMARANTHUS FROM EASTERN MEXICO AND SOUTHMOST TEXAS

During initial observations of Mexican Amaranthus, a series of collections
distinguished by inflated turret-like style bases and short, slender, basally
connate, erect, well-separated, spatulate sepals, and bracts much shorter than
the sepals were marked as possibly new. In Standley (1917), the taxon keyed
closest to A. brandegei Standl., a species known only from a few collections
in eastern Mexico (the type caliccuions is from Sinaloa, México), with which
it shared the characters of 3 stamens per male flower and short pistillate
floral bracts. A loan of the type collection of A. brandegei from UC confirmed
that our material is indeed is similar to A. brandegei, but A. brandegei actu-
ally has 5(—4) stamens per male flower, as previously noted by Sauer (1950),
not 2—3 as stated by Standley (1917). In its small perianth and short bracts
the new species also showed similarities with A. dubius Mart. ex Thell., which
also has 5 stamens per staminate flower.

Amaranthus dubius is a tropical, sometimes temperate, New World weedy
species common from the Caribbean region, southern Mexico to Panama
and northern South America. It is adventive in Africa and tropical Asia and
cultivated in Europe (Aellen 1959), Jamaica (Correll and Correll 1982), and
elsewhere. It has been reported as one of the very few polyploid (2n=64)
species in the genus (Grant 1959). Floral differences between A. brandegei
and A. dubius are illustrated in Sauer (1950); pistillate Howers of A. dubius
are illustrated in Aellen (1959), and both taxa are described in Standley
(1917). In the material seen, both A. brandegei and A. dubius are more ro-
bust plants with larger, leaves to 12 cm long and 8 cm wide with distinct
terminal inflorescences. I have seen additional material of A. brandegei from
Tom Van Devender’s recent collections from Sonora and Sinaloa, México,
which have larger leaves than the type

In A. brandegez the larger pistillate sepals are reflexed, thickened at the

800 Sipa 183)

base at maturity and the midveins often branch distally, the fruit wall is
smooth below the line of dehiscence but roughened above the line (Fig. 4F).
In contrast in A. dubius, all pistillate sepals are erect, they do not thicken at
the base at maturity and have distinct single green midveins bordered with
thin membranous margins and the fruit walls are typically elliptical and
often smooth throughout. In the new taxon (Figs. 4G, 8) the sepals are much
thicker, the margins are often not membranous medially, the sepals are dis-
tinctly thickened and join together at the base of the developing fruit and
are typically shorter than the developing fruit. They are somewhat contracted
in the lower half and slightly broader below the acute to obtuse-rounded
tips. The sepals are often green along the distal midvein and in the larger,
marginal sepals, the midveins are branched distally. The fruit wall is ir-
regularly rugose throughout and has a distinct smooth tower. Furthermore
the plants are much more slender, with small leaves that continue, though
diminished in size, well into the inflorescences—they do not develop dis-
tinct leafless terminal inflorescences as in the other two species. Also the

new taxon has only three stamens per male flower not 4—5 as in the other
species.
Amaranthus tamaulipensis Henrickson, sp. nov. (Figs. 4G, 5, 8). Typr:

MEXICO. Tamauttras: 13 mi N of Aldama on road to Soto la Marina, 25 Sep 1960,
M.C. Johnston & J. € epabfald 5698 (HOLOTYPE: TEX!).

A Amarantho brandevei-dubins differt calyce pistillato sepalis binis ad margines ovarit quam

<

Eelons altera majodau (vs. sepala abaxial ed hanes erectis (non distaliter reflexis
(vs. distaliter inflaci- ee ulato).

Erect to ascending, sometimes decumbent monoecious annuals 1.5—3(—
6) dm tall; stems branched at base, sparsely branched above or unbranched,
striate, glabrous, slender, tan-stramineous or suffused with betalin red, 1—
3.2 mm thick at base. Leaves largest near the base, reduced above, the peti-
oles 3—15(—21) mm long, one-half (when small) to as long as the leaf-blades;
leaf blades ovate, rhombic-ovate, (S—)10-—27 mm long, (2—)5—11(—18) mm
wide, tapering to an obtuse-rounded, typically emarginate tip topped with
the excurrent midvein, the base cuneate, decurrent along the petiole, the
margins rather crisped, white, the blades green above, gray-green with 3—
4 arcuate white, raised veins beneath. Flowers in glomerate axillary dicha-
sia from base of plant to the tip, either interrupted at the nodes, or crowded
or interrupted in terminal leafy spike-like inflorescences, the 2 lateral branches
of the dichasium unbranched, zig-zagged, to 4 mm long; bracts lance-ovate,
acute, conduplicate-cupped, green along midvein, otherwise scarious-mem-
branous, 0.9—1.2(—1.4) mm long, shorter than the sepals, persisting on
inflorescence axis where 0.4—0.8 mm apart; bracteoles 2, similar to the bracts
but 0.3—0.8 mm long, often membranous throughout, persistent or not;

HENRICKSON, Studies in New World Amaranthus 801

Fic. 8. Amaranthus tamautipensis. A. Terminal stem. B. Older stem showing inflorescence
development and leaves. C. Axillary inflorescence with most fruits gone showing persis-
tent bracts, He leaf. D. Mature tuberculate, dehiscent pericarp with calyx; note inflated
style base ate pericarp tip, seed. All from holotype (Aldama, Tamaulipas, Mex., Johnston
& Cue TEX, except C from Cameron Co., Tex., Runyon 2170, TEX). Magni-
fications as indicated.

staminate flowers: basal and scattered on the lateral dichasial shoots, sepals
4—5, oblong-ovate, acute-acuminate, 1.2—1.6 mm long, green along the excurrent
midvein, otherwise scarious; stamens 3(—4); pistillate flowers: sepals 5, separate
and not overlapping except where expanded and joined at the base, oblong-
spatulate, subequal with the two marginal sepals usually larger (broader)
than the other three, 1.1—1.6(—2.1) mm long, narrowed to 0.2—0.3(—4) mm
wide in mid portion, expanded, obtuse-rounded, to 0.25—0.6 mm wide near
the tip, rather scarious except along midvein, becoming gibbously thick-
ened, spongy and joined around ovary at base, the smaller 3 sepals 1-veined
and green along the vein above, the largest 2 sepals with the midvein green
and weakly branching distally (in some specimens the veins in all sepals
branching distally and the tips rounded to acute), the calyx shorter than
the mature fruit, the sepal margins typically sparsely glandular-puberulent.
Mature utricle obpyramidal, slightly biconvex, 1.5—1.7 mm long, to 1.1
mm wide, with age becoming brownish, rugose tuberculate throughout the

802 Sipa 18(3)

body with ridge-like branched folds, medially circumscissile but no specific
line of dehiscence evident, the ucricle tipped by a distinct, lighter tan, smooth,
rather inflated, dome-like style base 0.3—0.5 mm long, 0.5—0.6 mm wide,
with 3 broadly conic lobes at tip each bearing a papillate stigmatic line
along the inner margin that extends onto a slender 0.2—0.3 mm long tip.
Seeds black to reddish-brown, smooth, shiny, lenticular, orbicular, 1.0—1.2
mm wide, 0.6 mm thick, margins edged, slightly notched at base.

Additional collections: U.S.A. TEXAS. Cameron Co.: 2 mi S San Benito, 4 Jul 1939,
Runyon 2170 (TEX); E of Bena , 28 Oct 1927, Rose & Russell 24274 (US). MEXICO.
San Luis Potosi: Vicinity of El Salto above El Naranjo, 6 Apr 1960, Duke M3760 (NY).
Tabasco: Arrozal experimental del Depto. de Agronomia, Colegio Superior de Agricultura
ia |, km 22 carretera Cardenas-C gee 14 Apr 1980, Cowan 2913 (NY). Tamaulipas:
2 mi NE of Altamira, roadside weed, 1961, King 4056 (KF, NY, TEX, US-a more
eee nt plant with slender stems); 2 mi - Casas on new Victoria-Soto la Marina Hwy.,
28 Sep 1960, Johnston & Crutchfield 5770 (TEX).

As noted above, the species structurally is similar to A. brandeger and A.
dubius but is immediately distinguished from A. brandegez by its erect, not
reflexed calyx and from A. dubius by: 1) the narrow subequal, well-sepa-
rated, firm (not membranous margined) spatulate, acute to round-tipped
pistillate sepals that are shorter than the fruit and are expanded and united
at the base; 2) the distinctive roughened (not smooth) utricle wall with its
smooth, inflated, turban-like style bases; and 3); the 3 stamens. The sepals
along the edges of the compressed fruit are usually largest and typically
have veins branched below the tip. In A. dwbins the bracts and pistillate
sepals are larger, more elliptical, more membranous, stamens number 5(—

4) in male flowers and the utricle wall more gradually expanded Amaranthus
dubius, however, exhibits variability in bract length, sepal size, mature pericarp
configuration and may be confused with unarmed individuals of A. spinosus
L. with which it hybridizes (Grant 1959). Three varieties were described
by Moquin-Tandon (1848) under A. fristis L. and transferred to A. dubius
by Thellung (1914) (see Thellung 1914; Aellen 1959), but these are sel-
dom accepted. They involve differences in inflorescence development, color
and thickness.
A NEW TAXON FROM ARIZONA

Dr. Tom Van Devender requested identification of a series of Amaranthus
collections from the Tucson Mountains and Organ Pipe National Monu-
ment west of Tucson, Arizona. The collections show a number of character-

istics unlike any other southwestern Amaranthus. In Standley’s (1917) key,
the material, which has non-spatulate pistillate sepals, dehiscent utricles, a
terminal inflorescence, no spines, bracts shorter or equal to the pistillate
sepals, and the fruit shorter chan the sepals, keys to the aforementioned Amaranthus

HENRICKSON, Studies in New World Amaranthus 803

dubius, a species with much shorter sepals, smaller seeds, etc. The Arizona
specimens are small, erect, branched, monoecious annuals with coarse ovate
leaves with dichasial cymes extending from the mid-leaf axils into a leafless
terminal inflorescence. Staminate flowers have 5 sepals and 5 anthers; pis-
tillate flowers have broad-based, acuminate bracts shorter than or about as
long as the pistillate sepals (Fig. 9). The pistillate sepals are oblong, slightly
spatulate, but not clawed; the outer sepals are usually acute, somewhat spine
tipped, the inner ones are more obtuse or rounded. The sepals are distinc-
tive in being green throughout most of the surface except for the moder-
ately narrow scarious margins. As the sepals mature and dry, they reflex and
become thick and rigid. The fruit is dehiscent, about as long as the sepals,
with 3-2 styles, and is inflated and roughened above the line of dehiscence
(Fig. 9C),

While this appears to be an unremarkable assortment of characters, the
plants do not fall into any recognized taxon. Evidence of its possible rela-
tionship came from a specimen from Chiapas {32 km. N. Ocozocoautla along
rd. to Mal Paso, 19 Oct 1965, Breedlove G Raven 13572 (LL)} annotated by
J.D. Sauer as Amaranthus hybridus L._—atypical. This specimen has an inflorescence
arrangement characteristic of A. Aybridus, but the pistillate lowers have small
bracts and indurated sepals longer than the utricles—very similar to the

Arizonan specimens. However, the Arizonan specimens differ in having axillary
dichasia and terminate in a single spike of dichasia, an inflorescence type
more characteristic of Amaranthus species with clawed spatulate pistillate
flower sepals, e.g. A. obcordatus (Gray) Standley, A. chihuahuensis S.Wats, A.
torreyt (Gray) S. Wats. Somewhat similar plants have been found near Alamos
in southern Sonora, México. They are similar in development of the pistil-

late sepals, but in each the bracts are as long as or longer than the sepals
and one collection, more reflexed. Amaranthus hybridus differs from the new
taxon in the overall structure of the inflorescence, it produces a tight series
of slender, ascending-spreading shoots, the slender, longer bracts that well
exceed the flowers, and the membranous pistillate sepals that are typically
green only along the midvein.

A question arises as to the origin of this taxon. Is it just an obscure taxon
that has not been previously collected or is it a taxon of recent origin, per-
haps a product of hybridization? A brief analysis of pollen stainability of
the specimens revealed that pollen in Arizonan specimens varied greatly in
stainability with Cotton-Blue in Lactophenol (Radford et al. 1974), with
one collection having 99 percent stainability and other collections having
96, 89, 81,51, and 47 percent stainability. The two collections from Sonora,
both had high pollen stainability (97 and 95 percent). My conclusion is
that the taxon may be of hybrid origin, perhaps with A. Aydridus being one

804 Sipa 18(3)

parent, the other parent unknown. The only other species known from the
Tucson Mountains are A. fimbriatus (Torr.) Benth. and A. palmeri S.Wats.
(Rondeau et al. 1996), which are not parental candidates based on their
characteristics.

The Sonoran specimens both differ from the Arizona specimens. In Van
Devender 93-386 the pistillate sepals show green mostly along distinct sec-
ondary veins and subtending bracts have straight, long slender tips and a
membranous ovate base. The other specimen (Van Devender 92-1121) has
greenish sepals similar to the Arizonan specimens, but the subtending bracts
are as long as the sepals, strong reflexed below the tip, and are sharp pointed.

oe x tucsonensis Henrickson, nothosp. nov. (Figs. 5 » 9). Tyr:
I.S.A. ARIZONA: Pima Co.: Tucson Mts., crest of Gates Pass, W of Tucson,
14S,R12E,S10,SW 1/4, 3250 ft, 16 Sep 1988, TR. Van Devender 88-676 (HOLO-

type: ARIZ; isorypes: to be distributed)

—

A Amarantho hybridus differt an multo majoribus firmioribus viridibus scariosi-marginatis
(non omnino tenuis scariosus), bracteis floralibus ene sepalis piscillatis asanrile

vel Bley oenusnem | Bae 2 0 rlok longirib axe infloresce
glandulosi-c¢ glanduloso) et ante as 1.2—1.4 mm in diametro
(n

non 1.0 mm in diametro).

Erect, taprooted, mostly single-stcemmed or upwardly branched monoe-
cious annuals to 1.5 dm tall; stems yellowish, sometimes tinged with red,
striate, sparsely puberulent-pilose with crinkled, obscurely gland-tipped
moniliform hairs to 0.3(—0.6) mm long, glabrate. Leaves alternate; petioles
yellowish, 8-32 mm long; leaf blades rhombic to ovate, (1 5 )20=3 5745)
mm long, (7—)10—22(—28) mm wide, obtuse to rounded, often mucronate
at the tip, broadly cuneate with the margins narrowly decurrent along the
distal petiole at the base, the margins crisped, the lamina grayish-green,
glabrous to sparsely puberulent-pilose with crinkled, gland-tipped hairs to
0.3 mm long, the veins yellowish, impressed above, raised beneath. Flow-
dichasial cymes to 10 mm long, these forming

en

ers borne in axillary, compounc
discontinuous glomerules below, but coalescing into terminal, continuous
inflorescences 6—15 cm long, 8—10(—16) mm wide, the basal (oldest) flower(s)
staminate, developing lateral, sinuous, occasionally branching axes to | mm
thick, covered with the persistent bracts and bracteoles that subtended the
early deciduous male flowers. Staminate flowers: bracts broadly ovate at base,
acuminate above with the midvein excurrent, 1.7—2.5 mm long, scarious
except along the green midrib; sepals oblong-elliptical to lanceolate, 1.6—
2.2 mm long, 0.6-1.0 mm wide, acute-acuminate, scarious except along
the short excurrent midrib; stamens 5; filaments 1—1.8 mm long; anthers
0.8-1. (mm long, yellow. Pistillate lowers: bracts lanceolate to basally ovate,
1.5—3.5 mm long, 0.8—1.4 mm wide, ee shorter than the sepals, broadly
scarious below, the green midrib zone 0.2—0.5 mm wide, the midvein ex-

HENRICKSON, Studies in New World Amaranthus 805

Fic. 9. Amaranthus x tucsonensis. A. Growth habit. B. Mature calyx showing short subtend-
ing bract. C. Calyx and base of pericarp, tip of pericarp and seed. All from holotype (Pima
Co., Ariz. Van Devender 88-676, ARIZ). Magnifications as indicated.

806 Sipa 18(3)

current for 0.5—0.6 mm, bracteoles 2, similar but only 14 2.2 mm long;
sepals 5, oblong, oblong-spatulate to oblong-elliptical, 2.2— eee long,
().6—-1.2 mm wide, obtuse, rounded to emarginate, the outer ones more acuminate
with the midvein excurrent 0.1—0.3 mm at the tip, entire to erose distally,
green for 0.5—0.9 mm across the back, with scarious margins 0.15—0.2 mm
wide, glabrous to sparsely puberulent-pilose with crinkled, obscurely gland-
tipped hairs, unequal in size with the outermost sepal larger and more acute
than the inner round-tipped sepals, at maturity all sepals strongly reflexing,
Shawl and uniting at the base and overall becoming indurate. Fruit
2,2-2.5 mm long, circumscissile, the base smooth, the top roughened, inflated
re the zone of dehiscence, the 3 style bases with a U-shaped sinus; stig-
matic surfaces 1.2—1.3 mm long, puberulent with moniliform hairs. Seeds
broadly lenticular, slightly ovate, 1.2-1.4 mm wide, shiny black, with a
slight marginal rim.

Additional collections. U.S.A. ARIZONA. Pima Co.: ave le 1 Wash, Saguaro National
Monument, Tucson Mts., T13S,R11E,S35,NE 1/4, 2680 ft, - ae 1989, Bertelson 89-555
(ARIZ): Tucson Mts., crest of Gates Pass, T148,R12E,S1LO,SW 1/4, 3250 ft, 23 Sep 1988,
Van Devender 88-722 (ARIZ); Tucson Mts., crest near Radio nian NW of Trail’s End
=“ T14S,R12E,S4,NW 1/4, 4350 ft, 18 Sep 1988, Van Devender et al, 88-693 (ARIZ).

XICO Hee Rancho La Junta, 3.0 km NNE of Burapaco, 27°35'15"N, 108°52°W,

20m, 17 Mar 1993, Van ea et al. 93-386 (ARIZ); Mesa Masiaca, 6.5 km WNW of

san José de Masiaca, 26°46. 09°17.9 W, 200 m, 8 Oct 1992, Van Devender et al. 92-
1121 (ARIZ).

In the Tucson Mountains, west of Tucson, Arizona, the taxon occurs on
= slopes in desert scrub and grassland from 790-1430 m elevation (Rondeau
et al. 1996). In northern Sonora it is known from elevations about 200 m
in fields and roadsides. Flowering occurs from September to November or

Ss FS

after rains.
ACKNOWLEDGMENTS

I thank EF, GH, NY, TEX-LL, UC, US for loan of specimens, Charles Jarvis
(BM) for information regarding the type of A. polygonoides, U. Braun (HAL)
for information on the type of A. crassipes, Bobbi Angel (NY) for the line
drawings, Thomas Wendt for comments on the manuscript and the Span-
ish translation of the abstract, Guy Nesom for the Latin translations, and
the Plant Resources Center at the University of Texas, Austin for use of
facilities.

REFERENCES

AELLEN, P. 1959. aaa In: Hegi, G., Iluserierte flora von Mitteleuropa. 2nd ed. 3(2): 1605—

516. (Republished 19
Correit, D.S. and H.B. C ORRELL. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz.

—

HENrRICKSON, Studies in New World Amaranthus 807

CorreLt, D.S. and M.C. Jounsron. 1970. Manual of the vascular plants of Texas. Texas
Research Foundation., Renner.
Fawcett, W. and A.B. RENDLE. 1914. Amarantus L. Flora of Jamaica. 3:129—132.
JOHNSTON, I.M. 1944. Plants of Coens eastern Chihuahua, and adjoining Zacatecas anc
Durango, — ee Arbor. 25:133-182.
GRANT, W.F. 1 . Cytogenetic es in Amaranthus II. Natural interspecific es
ization ae Manabe dubius and A. spinosus. Canad. J. Bot. 37:1063-10
Gray, A. 1862. Enumeration of a collection of dried plants made by L. J. ee at Cape
San Lucas, &c. in Lower California, between AUEUSS, 1859 and ae 1860, and
communicated to the Smithsonian Institution. Proc. Amer. Acad. Arts. 5:153—17
Hoizincer, J.M. 1892. On Amaranthus crassipes. Bot. Gaz. 17254256, - XVii.
Raprorb, A.E., W.C. Dickson, J.R. Massry, and C.R. Bett. 1974. Vascular plant system-
atics. Harper & Row, New York. P. 219
Ronpbeau R.J., T.R. VAN DeveNnper, C.D. BerreLsen., P.D. JENKINS, R.K. Winson, and M.A.
Dimmitr. 1996. Annotated flora and vegetation of the Tucson Mountains, Pima County,
Arizona. Desert Pl. 12:3—46.
Sauer, J.D. 1950. The grain abiatenihs: A survey of their history and classification. Ann.
Missouri Bot. Gard. 37:561—
STANDLEY, P.C. ice ee L., N. Amer. FI. 21(2):101-119.
a A. 1912. Amaratacées. Tie = flore adventice de Montipellier. Mém. Soc. Nat.
. Nat. Math Cherbourg. 38: an os
ne 1914, Amarantus. In: Ascherson, P. and Graebner, P. Synopsis der Mitteleuropaischen
flora, 5(1):225- 356, ee 1919).
ULINE E.B. and W.L. Bray. 1895. A aoe aces of the North American species of
Amaranthus. Bot. Gaz. 19:267—272, 313-32

INDEX TO SCIENTIFIC NAMES

Amaranthus palmeri Blitum
ulbus poly gonioides polygonoides
berlandieri scleropoides Euxolus
brandegei tamaulepensis rassipes
chihuahuensis x texensis Roemeria
crassipes torryi polygonoides
Var. Crassipes tucsonensis Sarratia
var warnockil venulosus aa
dubius verticillatus Scler
briatus warnockii ania
graecizans Amarantus crassipe
hybridus Amblogyne
obcordatus polygonoides

808 Sipa 18(3)
BOOK RECEIVED

OLson, RicHARD K. and Tuomas A. Lyson, eds. 1999. Under the Blade.

The Conversion of Agricultural Landscapes. (ISBN 0-813 3-3597-

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Boulder, CO 80301-2877. $25.00 (pbk). 459 Pp. 15 Figures.

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Foreward Fauquier County, Virginia
Pretace and Acknowledgments Albemarle County, Virginia
About the Contributors Tillery, North Carolina
Introduction Lake County, Floride
Farmland Loss in America Ohio
A Landscape Perspective on Farmland DeKalb County, Hlinois
Conservation Waukesha County, Wisconsin
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Preserving Community Agriculture ina Lincoln, Nebraska (Antelope Commons
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Ethics and Aesthetics in the Loss of Farm- Northwest Arkansas
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A National Policy for Farmland Preser- Rocky Mountain region
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Case Studies Oregon
Swanton, Vermont rue County, Californ
Tompkins County, New York The Oxnard Plain, Califo ornia
Southeast Pennsylvania a

Sipa 183): 808. 1999

TWO NEW SPECIES OF TRIPOGON (POACEAE)
FROM INDIA

A.K. PRADEEP

Department of Botany
University of Calicut
Kerala 673 635, INDIA

C.N. SUNIL

Department of Botany
S.N.M. College, Maliankara
Kerala 683 516, INDIA

ABSTRACT

Two new species of Poaceae, Tripogon sivarajanii Sunil and T. vellarianus Pradeep from
the Western Ghats of Kerala, India are described and illustrated.

RESUMEN

Se describen e ilustran dos nuevas especies de Poaceae, Tripogon sivarajanii Sunil y T.
Pn Pradeep de los Western Ghats de Kerala, India.

The genus Tripogon Roem. & Schult., with about 31 species, is distrib-
uted in the Old World tropics (Mabberley 1997). In India, the genus has
ca. 12 species most of them occurring at an elevation above 900 m. Four
species, T. bromoides Roem. & Schult., T. narayanii Sreekumar et al., T.
ananthaswamianus Sreekumar et al. and T: capitellatus Jaub. & Spach. are known
from Kerala (Sreekumar & Nair 1991). During the course of floristic inves-
tigation on the Western Ghats of Kerala, the authors came across two in-
teresting specimens of Trzpogon which do not correspond to any of the pre-
viously described taxa. We conclude that they represent two new species
and we describe them here.

Tripogon sivarajanii Sunil, sp. nov. (Fig. 1). Tyee: INDIA. Kerata. Idukki De.

Valakettimala near Moolamattam, 1020 m, 2 Sep 1997, C.N. Sunil 2117 (HOLOTYPE:
MH; tsoryres: BRIT, K, L, PBL).

Tripogoni trifido affinis, T-stvarajanii ligulis glabris, lemmatis aristis 3—3.5 mm longis,
palea apicaliter 2-lobata facile distinguendus.

Tufted perennial herb. Culms 40-70 cm high; nodes glabrous. Leaves
with the blades 15—40 cm long and 2—3 mm broad, linear, acuminate, villous

Sipa 18(3): 809-814. 1999

810 Sipa 18(3)

Fic. |. Tripogon sivarajanii Sunil. A. Habit. B. Spikelet. C. Lower glume. D. Upper glume.
E. Lemma. F. Palea. G. Stamens and Pistil. A-G, drawn from holotype.

PRADEEP AND SUNIL, Two new species of Tripogon 811

on the upper surface with short dense hairs intermingled with long scat-
tered shaggy hairs, glabrous below, the midrib inconspicuous; ligule a fine,
glabrous membrane; sheath smooth. Racemes 20—45 cm long, with 25—60
spikelets; rachis smooth, glabrous. Spikelets 5—8-flowered, 7-8 mm long;
callus hairy. Lower glume ca. 2 mm, lanceolate, shallowly notched on one
side. Upper glume 3-4 mm long, lanceolate, acute or acuminate at the apex.
Lemma 3—3.5 mm long, 3-nerved and 1-awned, glabrous, 2-lobed at the
apex, the lobes acuminate, awned in the sinus; awns 3—3.5 mm, more or
less equal to lemma, scabrid. Palea 2-4 mm, elliptic, 2-lobed at the apex,
2-keeled, winged, the wings narrow, the margins ciliate. Stamens 3; an-
thers ca. 1 mm, yellow tinged with pink. Ovary 0.25 mm, globose; styles
2; stigma feathery, purple. Grains unknown.

Distribution.—Tripogon sivarajanii is endemic to Kerala and is known only
from the type locality, Valakettimala near Moolamattam in Idukki District
of Kerala.

Ecology. —This species grows from 1000-1300 m elevation along grassy
hill slopes and flowers from September to December.

Tripogon sivarajanii closely resembles, T: trifidus Munro ex Stapf, a spe-
cies distributed in the eastern part of India in having 20-45 cm long racemes,
5—8-flowered spikelets, laterally lobed and notched lower glumes and the
awns of the lemma being neither geniculate nor twisted. The two species
can be separated as follows:

1. Ligules ciliate, densely on the margins towards the apex; upper glume 2-

toothed with a short awn at the apex; lemma bearded at base, 2-lobed at

apex, awned in the sinus, the lobes awned at the apex; median awn twice as

long as lemma; palea obtuse at the apex T. crifidus
. Ligules glabrous throughout; upper glume never 2-toothed or awned at apex;

lemma glabrous at base, 2-lobed at apex, awned in the sinus, the lobes never

awned at the apex, median awn as long as or shorther than lemma; palea

2-lobed at the apex ‘T. sivarajanii

—

The specific epithet honors the late V.V. Sivarajan, one of the foremost
authorities on the flowering plants of Kerala. Asa staff member of the University
of Calicut, he had published over 25 new species, thus the dedication of
this new Poaceae to him is appropriate and well-deserved.

Tripogon vellarianus Pradeep, sp. nov. (Fig. 2). Tyee: INDIA. Kerata. Kozhikode
Dt.: Vellarimala, 11°25.877'N, 76° 06.765’E, 1300 m, 17 Oct 1997 (fl), A.K. Pradeep
56110 (HoLoTyPE: MH; isotypes: BRIT, K, L, PBL).

Tripogon vellarianus sine cognato propinquo a congeneris aliis habitu valde caespitoso,
foliis 8-15 mm latis, spiculis 10-17 mm longis, lemmate 2-lobato 3-aristato vel 1-aristato,
aristis lateralibus minutis ad nullis distinguendus.

812

Sipa 184)

AN

Ne
JOAN

=
Z

Ss
Z

|

2
Fic. 2. Tripogon vellarianus Pradeep. A. Habit. B. Spikelec. C. Upper glume. D.
E. Lemma. FE. Palea. G. Lodicule. A—G, drawn from holotype.

Lower glume.

PRADEEP AND SUNIL, Two new species of Tripogon 813

Tufted perennial herb. Culms 30-90 cm tall, erect; nodes glabrous. Leaf
blade 30-60 cm long, 8-15 mm wide, linear, base of blade glabrous. Sheaths
closely clasping, rigid, glabrous; ligules indistinct. Racemes 30-40 cm long,
rachis stout, glabrous. Spikelets 10-17 mm long, distant, dorsiventrally
flattened, 8—10-flowered; callus bearded. Lower glumes 4 3 1.25 mm, lan-
ceolate, acuminate at the apex, purplish tinged dorsally, glabrous. Upper
glumes 5 3 1.5 mm, elliptic-lanceolate, acute at the apex, prominently 3-
nerved, glabrous. Lemmas (excluding the awn) 5 3 2 mm, ovate-lanceolate,
3-nerved, median awn 3 mm long, lateral awns absent or up to 5 mm long
often closely adpressed to the median awn, glabrous. Palea 4 3 2 mm, hya-
line, narrowly elliptic, keeled and winged, keels minutely puberulous. Lo-
dicules 2, ca 0.25 mm, quadrate, apex coarsely 3-toothed. Stamens 3; an-
thers 1.5 mm long, oblong; filaments 0.7 mm long, slender, glabrous. Ovary
0.25 mm, obovate; styles 2, slender, hyaline, 0.8 mm long; stigmas feath-
ery, purplish, 0.6 mm long. Grains not seen.

Distribution.—Tripogon vellarianus is known only from Vellarimala on the
Western Ghats of Kerala in India. This species is to be expected from adja-
cent hill ranges of Wayanad and Kannur districts.

Ecology.—This species occurs from 1000-1500 m in wet rocky grasslands
adjoining evergreen forests. It flowers from October to January.

Tripogon vellarianus is a distinctive species characterised by its strongly
tussock-forming habit, broad leaves and large spikelets. Thomas A. Cope
(Kew, pers. comm.), after examining our material, commented “in major-
ity of the lemmas the lateral awns are minute or absent altogether. I cannot
recall another species in the genus that is so reluctant to produce lateral
awns.” It does not compare with any of the species so far known under the
genus, although its entire lemma sometimes recalls that of Tripogon major
Hook. f., a species widespread in tropical Africa. Dr. P.V. Sreekumar, Port
Blair (pers. comm.) wrote “I find it extremely difficult to accommodate even
under Tripogon; and I believe lemma is continuous with the awns.” How-
ever, apart from the entire lemma, in every other respect it corresponds exactly
with the genus Tripogon.

ACKNOWLEDGMENTS

We thank P.V. Sreekumar, Botanical Survey of India, Port Blair and Thomas
A. Cope, Kew for useful comments on our materials. We also thank J.F.
Veldkamp, Rijksherbarium, Leiden for various help. The work was sup-
ported by a grant from the Department of Science and Technology, New
Delhi, ref. no. HR/SY/B-05/96.

814 Sipa 18(3)
REFERENCES

Masserey, D.J. 1997. The plant-book. Ed. 2, Cambridge University Press.
SREEKUMAR, P.V. and V.J. Nair, 1991. Flora of Kerala
Calcutta.

Grasses. Botanical Survey of India.

BARNEBYDENDRON, A NEW GENERIC
NAME (FABACEAE, CAESALPINIOIDEAE,
DETARIEAE, BROW NEA GROUP)

JOSEPH H. KIRKBRIDE, JR.
USDA, Agricultural Research Service
Systematic Botany and Mycology eeu

Room 304, Building O11A, BARC-W
alee Me 2070522550, US; AL

" LVS ida -LOV

ad!

ABSTRACT

w generic name, Barnehydendron, is proposed for Phyllocarpus Riedel ex Tul. which is
an illegitimate, later homonym of Phy/locarpus Riedel ex Endl.

RESUMEN
Se propone el nuevo nombre genérico, Barnebydendron, para reemplazar a Phy/locarpus

Riedel ex Tul., ya que, este nombre es ilegitimo y un homénimo posterior de Phy/locarpus
Riedel ex En dl.

RESUMO

O novo nome genérico, Barnebydendron, é proposto para substitutir a Phy//ocarpus Riedel ex
Tul., visto que este nome é ilegitimo e homénimo posterior de Phy//ocarpus Riedel ex Endl.

In 1842 Endlicher (1842) presented the genus Phyllocarpus Riedel ex Endl.
on page 97 of the second supplement to his Generum plantarum:

6720/1 Phyllocarpus RID.

Lonchocarpus pterocarpus DC. Prodr. I. 260. typum distincti generis (Phyllocarpus
Riedel msc.), a Platymiscio et Miscolobio (Gen. pl. n. 6719 et 6720) leguminis membranacei
sutura vexillari alata diversi, constituit.

Endlicher validly published Phy//ocarpus with a single element, L. pterocarpus,

which is automatically the type of the genus. He distinguished it from

Platymiscium Vogel and Miscolobium Vogel (= Dalbergia L. f.) by its membra-

nous legume with the vexillary suture variously winged. Lonchocarpus pterocarpus

DC. (=Deguelia scandens Aubl.) has winged fruits that are somewhat fragile

(MacBride 1943), and so the diagnosis could well describe material of L.

pterocarpus.

n 1843 Tulasne (1843) described Phyllocarpus Riedel ex Tul., specifi-

cally excluding Endlicher’s Phy/locarpus, “non Endl. Gen. Supp I, p. 97, n.

6720.” He also described P. riedelii Tul. with a short description using no.

Sipa 18(3): 815-818. 1999

816 Sipa 18(3)

1022 in Guillemin’s Brazilian herbarium, and since it was the only species
in the genus, it is the type of Phy//ocarpus Riedel ex Tul. In 1844 Tulasne
(1844) republished his generic description verbatim, amplified the description
of P. riedel7i, discussed the generic relations of Phyllocarpus, and illustrated
the species. He also stated that specimen no. 1022 in the Guillemin her-
barium was a Riedel collection. Apparently Tulasne examined Riedel’s col-
ection in Guillemin’s herbarium, and noted that Endlicher’s Phy//locarpus
did not correspond to what Riedel had intended. Therefore he redescribed
Phyllocarpus using the material of Riedel, and thereby created a second Phy/locarpus
which is a later, illegitimate homonym.

Geesink (1984) separated Deguelia Aubl. from Derris Lour., and p
all the American species of Derris in Deguelia, including L. pterocarpus. He
cited Endlicher’s Phy//ocarpus as a later synonym of Deguelia. The type of
Endlicher’s Phy//ocarpus is now a later synonym of Deguelia scandens with

—

—

aced

—_

the following synonymy:

Deguelia scandens Aubl., Hist. pl. Guiane 2:750, t. 300. 1775.
Lonchocarpus pterocarpus DC., Prodr. 2:260. 1825.

Derris guianensis Benth., J. Linn. Soc., Bot., 4, suppl.: 106. 1860.

Derris scandens (Aubl.) Pittier, Contr. U.S. Natl. Herb. 20:41. 1917, non
Derris scandens Benth., 1860.

Derris pterocarpa (DC.) Killip, J. Wash. Acad. Sci. 26(9):360. 1936.

This species is used as a fish poison (Killip 1936; Uphof 1968) in northern
South America.

In 1912 Tulasne’s Phy/locarpus was discovered in Guatemala, and John
Donnell Smith (1913) mistakenly described it as a new species, P. seprentrionalis
Donn. Sm. It was later discovered as far south as Panama (Standley & Steyermark
1946; Woodson & Schery 1951; Holdridge & Poveda 1975). Barneby (1996)
reviewed the genus, and concluded that it is monotypic and that P. septentrionalzs
is a later synonym of. P. riede/ii.

Phyllocarpus in the sense of Tulasne has traditionally been accepted as a
genus in the Fabaceae. It has been used in systems of legume classification
(Bentham 1865; Hutchinson 1964; Cowan & Polhill 1981; Polhill 1994),
floras (Bentham 1870; Britton & Rose 1930; Standley & Steyermark 1946;
Woodson & Schery 1951; Holdridge & Poveda 1975; Isely 1975), and
morphological studies (Gunn 1991). From the descriptions in these works,
it is obvious that the authors mentioned above were referring to Phy/locarpus
Riedel ex Tul. However, in Britton and Rose (1930) and Woodson and Schery
(195 1) its place of publication was given as the second supplement to Endlicher’s
Generum plantarum (1842), and also in Britton and Rose (1930) its type was
cited as “Phy/locarpus Pterocarpus Riedel,” an unpublished epithet.

KiRKBRIDE, JR., Barnebydendron, a new generic name 817

Isely (1975) was the first to note that Tulasne’s Phy//ocarpus was a later
homonym of Endlicher’s PAy/locarpus, and therefore illegitimate. Barneby
(1996) also recognized that Tulasne’s Phy//ocarpus was illegitimate. There-
fore the following new name is presented for Tulasne’s Phy/locarpus:

Batnebydendron J.H. Kirkr., nom. nov. Phyllocarpus Riedel ex Tul.,
Nat. Bot., sér. 2, 20:142. 1843, non Phy/locarpus Riedel ex Endl., 1842. Type: ee
riedel1i Tul. (= Baeoids on riedelii (Tul.) J.H. Kirkbr.).

ear esee —— ae J. a ere Toe nov. seaaacie riedelii
. 1843. Typ : BRAZIL. Rio de Janeiro,
pees ie a pe ae ra rete No. 1022 Reet ay (HOLOTYPE: P-
; ISOTYPE: US!).

oe Septentrionalis Dona. Sm., Bot. Gaz. 55:433. 1913. Types: GUATEMALA.
Departamento Z Gualan, Feb 1912 (A), WP. Cockerell s.n. (LecroryPE, US861342!
(Standley and Sick 1946; labelled by Standley as “type”}; soLecrorypes, US1337584!
{fragments}, US1337585! {fragments]}); May 1912 (fr), E. Morris s.n. (PARATYPES, US] 1337586!
[fragments], US1337587!, US1337588!).

I take great pleasure in dedicating this genus to Rupert C. Barneby who
has contributed so much to legume systematics. During my doctoral stud-
ies, I shared an office with Rupert at the New York Botanical Garden, Bronx,
New York. Rupert taught me many things, and significantly contributed
to my botanical training.

In his review Barneby (1996) gave the distribution of B. riedelii as Cen-
tral America from Guatemala to Panama, the Purtis basin in the southwestern
Brazilian Amazon, and the vicinity of Rio de Janeiro. It has recently been
collected in Amazonian Peru (N. Begarzo 150, 20 May 1980 [US}) and Venezuela
(J. Lissot s.n., 24 Nov 1975 [US}) partially filling the gap between Central
America and the southwestern Brazilian Amazon. Barneby (1996) cast doubt
on whether it was native to the vicinity of Rio de Janeiro. As far as he knew,
it has not been collected as a wild plant near Rio de Janeiro.

Barnebydendron has been reported as cultivated in southern Florida, Cuba,
Guatemala, Costa Rica, and Panama. Wild trees are 15—35 m tall, and cul-
tivated flowering specimens are 5—12 m tall. It flowers in the dry season
without leaves, and is full of flowers that are described as various shades of
intense red. Standley and Steyermark (1946) stated, “in general appearance
and in the color of its blossoms this tree suggests poinciana (De/onix) and is
almost equally showy,” and Woodson and Schery (1951) wrote, “the abun-
dant scarlet flowers rivaling in showiness those of the Royal Poinciana (De/onix
regia).” This beautiful tree merits more widespread cultivation.

ACKNOWLEDGMENTS
thank Drs. L.J. Dorr (US), D.A. Neill (MO), A.Y. Rossman (BPI), and

818 Sipa 18Q)

J.H. Wiersema (BARC) for their reviews and constructive criticisms of the
manuscript and M. Cristina Garcia Kirkbride and Tarciso S. Filgueiras (IBGE)
for reviewing the Spanish and Portuguese abstracts.

REFERENCES
Barnesy, R.C. 1996. Neotropical Fabales at NY: asides and oversights. Brittonia 48:174—

187.
a G. 1865. Ordo LVII. Leguminosae. In: G. Bentham and J.D. Hooker, eds. Gen.
. 1(2):434—600. Lovell Reeve & Co., London, England.
bev G. 1870. XCV. Phyllocarpus Riedel n: C.EP. von Martius, ed. Fl. bras. 15(2):61-
62, t. 19.

ee TON, N.L. an ng) N. Rose. 1930. 9. Phyllocarpus Ried.,; Endl. Gen. Suppl. 2:97. 1842.
r. Fl, 23(4):219.

oo ne ar R.M. Pout. 1981. Tribe 4. Detarieae DE. (1825). In: R.M. Polhill and
P.H. Raven, eds. Advances in Legume Systematics 1:1 17-134. Royal Botanic Gardens,
Kew, age

ENDLICHER, 5.L. 1842. Gen. pl., suppl. 2. F eck, eee a

GEESINK, R. oi Seal Millettiearum. a ‘Bor Ser. on

GuNN, C.R. 1991. Fruits and seeds of — in the subfi an Caesalpinioideae (Fabaceae).
Techn Bull. U.S.D.A. 1755:1—40

Horprince, L.R. and L.J. Povepa A. os Arboles de Costa Rica, vol. 1. Centro Cientifico
Tropical, San José, Costa Rica.

HUTCHINSON, J. 1964, The genera of flowering plants, vol. |. Oxford University Press, London,

. d.

IsELy, 1975. Leguminosae of the United States: H. Subfamily Caesalpinioideae. Mem.
New om Bot. Gard. 25(2):1—228.

Kip, E.P. 1936. New plants mainly from western South America
Sci. 26:358-361.

Macpribe, J.F. 1943. Leguminosae. Bean family. = J.B Macbride, Flora of Peru. Field
Mus. Nat. Hisc., Bot. Ser. 1 3(part 3, no. 1):1—-50

PoLuitt, R.M. 1994, Complete synopsis of (eae genera. In: RA. Bisby, J. Buckingham,

B. Harborne, eds. Phytochemical dictionary of the Leguminosae, pp. xlix—lvii. Chapman
& Hall, Penden, England.
SmitH, J.D. 1913. Undescribed plants om Guatemala and other Central American re-
publics, XXXVI. Bot. Gaz. 55:431-43
STANDLEY, P.C. and J.A. STEYERMARK. 1946. Phila Riedel. In: P.C. Standley and J.A.
Steyermark, eds., Flora of Guatemala. Fieldiana, Bor. 24:144—-14

Tutasne, L.-R. 1843. Nova oo proponit genera in Leguminosarum classe. Ann. Sci.
Nat. Bot., sér. a 20:136-

TUL ASNE, L. -R. 8 44 Lé égumine uses arborescentes de l’ Amérique du Sud. Arch. Mus. Hist.
Nat. | ee

Upnor, J.C.T. 1968. Dictionary of economic plants, ed. 2. J. Cramer, Lehre, Germany.

Woopson, R.E. and R.W. ScHEry. 1951. 5. Phy ae Riedel. In: R.E. Woodson and R.W.
Schery, eds., Flora of Panama, part 5, fasc. 3, pp. _ Ann. Missouri Bot. Gard. 38:324—325.

V. J. Wash. Acad.

A NEW ENDEMIC SPECIES OF MENTZELIA SECT.
BARTONIA (LOASACEAE) FROM NEW MEXICO

THOMAS K. TODSEN
Department of Biology
New Mexico State University
Las Cruces, NM 88003, U.S.A.
ABSTRACT
Mentzelia conspicua T.K. Todsen, endemic to the Rio Chama basin of New Mexico, is
illustrated and described. It is compared to a similar species, Mentzelia laciniata (Gray) Darl.
The distribution of the two species in Rio Arriba County is shown
RESUMEN

Se ilustra y se describe Mentzelia conspicua T.K. Todsen, endémica de la cuenca del Rio
Chama en Nuevo México. Se compra con la especie semejante, Mentzelia laciniata (Gray)
Darl. Se muestra la distribucién de las dos especies en el condado de Rio Arr

Key Woros: Mentzelia, Loasaceae, New Mexico
Mentzelia conspicua T.K. Todsen, sp. nov. (Fig. 1). Type: U.S.A. New Mexico.

Rio Arriba Co.: W end of El Vado Dam in deep road cut on NM Hwy 112, T28N,
R2E, 19 Aug 1991, Todsen 97-8-11 (HOLOTYPE: NMC; tsorypr: LA).

Mentzeliae daeenifatae ai ) Dz url, Similis sed differt. Differentiae sunt: staminum non-
petaloideum; ongitudine variante, verticillum intimum

7-10mm, ein extimum Cre 30mm; paginae superae foliorum virides cum pilis dispersis.

Erect, strict, herbaceous perennial to 60 cm tall, pubescent with small
retrorsely barbed hairs and long barbed hairs with tips acute and bases sur-
rounded by support cells. Rosette leaves and cauline leaves pinnately lobed,
rachis 2—3 mm wide, lobes usually opposite, 6-15 mm long, 2—3 mm wide,
lower surfaces pubescent with stout curved, pointed hairs and fewer, smaller
retrorsely barbed hairs, upper surfaces sparsely pubescent with few scattered
pointed hairs. Bracts linear, usually entire but occasionally with 2 narrow
lobes at base. Flowers opening late afternoon. Calyx lobes 5, 8-12 mm long,
deltoid, acuminate, calyx tube 1.5—2 cm long; petals 10, in two whorls,
the inner whorl of 5 petaloid staminodes, ovate to lanceolate, golden yel-
low, glabrous, outer whorl 25-40 mm long, 8-11 mm wide, inner whorl
22—35 mm long, 7-10 mm wide. Fertile stamens numerous, filaments all
filiform, outermost whorl filaments 20—27 long, innermost 7—10 mm long,
anthers 1 mm long; pistil 1; style 1, 24-32 mm long; stigma 1.5 mm long,
at anthesis 9-12 mm above the anthers. Capsule cylindrical, 15-22 mm
long, topped by persistent calyx lobes. Seeds lenticular, dark gray to black

Siva 18(3): 819-822. 1999

820 Sipa 18(3)

Fic. 1. Mentzelta conspicua (Todsen 91-8-5). A. Plant. B. Leaf. C. Capsule. D. Flower.

when mature, 2—2.8 mm long, wing narrow, to 0.1 mm wide. Chromo-

some number n=10. Self compatible.
Distribution and ecology.—In the upper Rio Chama drainage of Rio Arriba

County, New Mexico (Fig. 2), 1800-2250 m, on road cuts and steep barren
hillsides of gray to red shales and clays of the Mancos and Chinle forma-

tions; Howering July to early October.

TODSEN, A new species of Mentzelia from New Mexico 821

Fic. 2. Distribution of Mentzelia conspicua (circles) and M. daciniata (triangles) in Rio Arriba

County, New Mexico.

Additional specimens: U.S.A. NEW MEXICO. Rio Arriba Co.: 8 mi S of Canjilon, 24
Jul 1928, Wolf 2908 (CAS, CH, RSA); El Vado Dam, the Point, 22 Jul 1949, Castetter and
Dittmer 6559 (UNM); El Vado Dam, 22 Jul 1949, Castetter and Dittmer 6565 (UNM); 22

mi S of Tierra Amarilla, 1954, Waterfal/ 11126 (BRIT, COLO); 2.2 mi S of Piste Aa
Museum, 21 Aug 1964, chromosome voucher n=10, Atsatt 517 (LA); US84, 3.1 miSo
turnoff to ee 21 Aug 1964, chormosome voucher n=10, Aftsatt 518 (LA , US); ee
84, 2.4 mi S of Ghost Ranch Museum, fine red soil, 17 Sep 1968, Thompson 3558 (LA),
US64, es mi E of Tierra Amarilla, 30 oF O74, Todsen s.n. ae Echo Amphitheater
Campground,, 19 Aug 1991, Todsen 91-8-1 (LA, NM C); US 84 1 S of entrance to Ghost
Ranch, 19 Aug 1 Ae Todsen 91-S8-2 (LA, NMC, NY); US84, 4.3 mi iS of jet, with US 64E,
Todsen 91-8-3 (N LA, NY); US84, 9 mi S of Cebolla, 19 Aug 1991, Todsen 91-8-4
(NMC); NM 95, just a of Rio Chama bridge, 19 Aug 1991, Todsen 91-8-5 (LA, NMC,
NY); NM 95, Heron Lake Park Visitor Center, 19 Aug 1991, Todsen 91-8-6 (NMC): Ghost
Ranch, Kitchen Mesa Trail, 8 Jul 1994, Stvrnsky and Lowery 2768 (NMC, UNM); NM 112
5 mi W of El Vado Dam, 18 Aug 1994, Todsen 94-8-3 (NMC); NM312, 3 mi E of USS84,

18 Aug 1994, Todsen 94-8-4 (NMC); NM95, NW end of El Vado L ee 18 Aug 1994,
Todsen 94-8-5 (NMC); Forest Road 151,S of Big Eddy Pullout, Chama River Canyon Wilderness,
6 Oct 1997, Todsen 97-10-1 (NMC). Torrance Co.: 16 mi E of Clines Corners, 17 Aug
1968, Waterfall 15166 (BRIT, OSU, RSA)

Thirty-plus years ago, H.J. Thompson (personal communication) noted
that Mentzelia specimens collected from the upper Rio Chama drainage differed
from typical Mentzelia laciniata as described by Darlington (Darlington 1934),
particularly in the presence of variable stamen lengths. He stated that the
only other Mentzelza showing this characteristic was M. crocea Kellogg (=

—

822 Sipa 184)

M. lindleyi Torr. & Gray), so far as he knew. He later distributed a tentative
description of what he called M. conspicua, but never published it.

Since all the earlier collections of Mentzelia from the upper Rio Chama
basin were identified as M. /aciniata, the differences between the latter spe-
cies and M. conspicua are emphasized. For M. conspicua, 1) leaf lobes are usu-
ally opposite vs. usually alternate; 2) upper leaf surfaces and capsules have
scattered pubescence vs. dense pubescence; 3) petaloid stamen filaments are
absent vs. present; 4) stamen lengths for an individual flower are variable
(7—22 mm) vs about three-fourths the length of the petals. Other more subjective
differences are that M. conspicua plants are stouter and taller and the flowers
are much larger (6—9 cm vs 3.5—4.5 cm).

There was an anomalous collection of M. conspicua from 16 mi E of Clines
Corners in Torrance County (Waterfall 15166) 160 km SE of the nearest
Rio Chama site. I have examined the specimens at BRIT and RSA and have
a photocopy of the specimen at OSU and they are indeed M. conspicua. Attsatt,
Thompson and I have independently searched the stated location and have
found neither specimens nor characteristic habitat. The original site was
along US Highway 66, which was completely redone to form Interstate Highway
40. It may be that the location was extirpated by that construction.

ACKNOWLEDGMENTS

Tam indebted to H.J. Thompson for his sharing research data and advice
on Metzelia conspicua as he called the taxon. thanks go to Linda Reeves
for her drawing and to friends and colleagues who have offered advice and
encouragement. Finally, my appreciation to the reviewers and the editor
for their cogent comments and criticisms.

REFERENCES

esi J. 1934. A monograph of the genus Mentze/ia. Ann. Missouri Bot. Gard. 21:103—
2:

NOMENCLATURAL NOVELTIES IN NUPHAR
(NYMPHAEACEAE)

DONALD J. PADGETT!
Department of Biology
Southwest Missouri State University
Springfeld, MO 65804, U.S.A
ABSTRACT

Recent revisionary and cladistic studies in Nuphar Sm. resulted in the following nomenclatural
tio cribed (sect. Nuphar and sect. Astylus), five new

amendments: two novel sec
combinations are proposed (N. i orbiculata, N. advena subsp. ozarkana, N. advena
subsp. wlvacea, N. pumila subsp. sinensis, N. pumila subsp. oguraensis), and one species, N.
oguraensis Miki, is lectotypified.

RESUMEN

Los estudios recientes, cladisticos y revisiones de Naphar Sm. dan por resultado las siguientes
enmiendas nomenclaturales: se debe ies secciones nuevas (sect. Nuphar y sect. Astylus),
proponen cinco combinaciones nuevas (N. advena subsp. orbiculata, N. advena subsp. ozarkana,
ee subsp. w/vacea, N. pumila subsp. sinensis, N. ae subsp. ogaraensis), y se lectotipifica

N. ogaraensis.

Nuphar Sm. (Nymphaeaceae) is a relatively common genus in north temperate
regions of North America, Europe and Asia. Inhabiting ponds, lakes, streams
and slow moving rivers, these perennial herbs are characterized by floating
and submersed leaves and solitary, globose, yellow flowers.

While conducting monographic research on Nuphar, morphological analyses
initially suggested that the genus was represented by two sets of species,

Old World and New World segregates (Padgett et al. 1996). Cladistic analyses
of representative internal transcribed ee (ITS) sequences (nr DNA) and
matK (cpDNA) sequences subsequently offered strong support for the same
two infrageneric lineages (Padgett 1997; Padgett et al., in press). Accord-
ingly, the classification of Nwphar is emended to include two sections.

Nupbhar sect. Nuphar is characterized by five sepals, anthers no more than
half the length of the filaments, and urceolate fruits with elongated necks,
or “styles” of some authors. It contains all Eurasian species with a single
species (N. microphylla (Pers.) Fern.) represented in northeastern North America.
Nuphar sect. Astylus is characterized by 6-12 sepals, anthers 1-2 times the

'Department of Biological Sciences, Bridgewater State College, Bridgewater, MA 02325,

U.S.A. dpadgett@bridgew.edu

Sipa 18(3): 823-826. 1999

824 Sipa 18(3)

length of the filaments, and ovoid fruits lacking apparent necks. It is en-
tirely North American in distribution.

Nuphar sect. Nuphar. Tyee: Nuphar lutea (L.) Sm. in Sibth. & Sm., Fl. Graec. Prodr.
1:361. 1808 (1809).
Nuphar sect. Astylus D. Padgett, sect. nov. Type: Nymphaea advena a es
kew. 2:226. 1789. Nuphar advena (Ait.) Ait. f., Hort. kew. ed. 2, 3:295.

luplo usque longiores; fructus ovoideus,

Sepala 6-12; antherae elongatae nt
haud infra discum constrictus; cus stigmaticus integer (rarius crenatus).

Sepals 6-12; anthers long, 1-2 times the length of the filaments, fruit
barrel-shaped without a prominent neck, stigmatic disk entire (rarely crenate).
North America. Type species: Nuphar advena (Ait.) Ait. f.

The synonymy of Nuphar is extensive and only recently has been assessed
from a world-wide perspective (Beal 1956; Padgett 1997). However, since the
primary purpose of this article is to make these new names available, com-
plete synonymy will be withheld for a later publication (or see Padgett 1997).
It should be noted, however, that Beal (1955; 1956) greatly broadened the
species concept in his revision of Nuphar, thus placing into synonymy man
previously recognized species under the type species N. /wtea (L.) Sm. While
information gleaned from the recent phylogenetic studies of the genus pre-

cludes any attempt to treat North American entities at a subspecific level
under N. /utea (as urged by Beal [1956}), an alternative classification does
adopt infraspecific taxa for some species (Padgett 1997). These taxa, as subspecies,
are based on groups of populations with distinctive characters which ex-
hibit geographical, and in some cases ecological, integrity but otherwise
all within the range of morphological variability for the species. A more
detailed account of studies that support the taxonomic decisions reflected
here will be forthcoming elsewhere (or see Padgett 1997).

As recognized by Padgett (1997), Nuphar advena sensu lato 1s a polymor-
phic species in section Asry/us that ranges throughout most of southeastern
North America. It is comprised of four recognizable, but morphologically
strongly overlapping and inter-grading subspecies.

The reduction in rank of the three taxa that follow is based on examina-
tion of numerous herbarium specimens of this species complex from throughout
its range, coupled with fieldwork, as well as multivariate statistical analy-
ses (Padgett, in prep.). Infraspecific ranking of most of these formerly rec-
ognized species has been supported in opinion by other workers (e.g., Yatskievych
& Turner 1990; Wiersema & Hellquist 1997).

a es ) Aic. f. subsp. orbiculata (Small) D. Padgett, comb.
. Bas Nymphaea orbiculata Small, Bull. Torrey Bot. Club 23:128. 1896.

Rane rae (Small) Standl., Field Mus. Nat. Hist. Bot. Ser. 8: aren 1931.
Nuphar lutea (L.) Sm. ae or ies (Small) E.O. Beal. J. Elisha Mitchell Sci. Soc.

Papcett, Nomenclatural novelties in Nuphar 825

72:338. 1956. “orbiculatum” Type: U.S.A. GEORGIA. paddies Co.: eons or near
the Ochlockonee River near Thomasville, Sma//5.n. (LECTOTYPE d by Miller

& Standley (1912): NY; IsoLecrorypes: F not seen, NY).
AUpag phic (Ait.) Ait. f. subsp. ulvacea (Mill. & Stand.) D. Padgett,
mb. nov. Basionym: Nymphaea eres Mill. & Standl., Contr. U.S. Natl. Herb.

es 97. or 5 Nuphar ilvaceum (Mill Standl.) Standl. ‘Field Mus. Nat. Hist. Bot.
Ser. 8:310—11. 1931. pee lutea io ) Smith subsp. ene (Mill. & Standl.) E.O.
Beal., J. Beha Mitchell Sci. Soc. 72:336. 1956. “avlvaceum’. Type: U.S.A. FLORIDA:

Blackwater River near oe ne 6409 (HOLOTYPE: US, not seen; isoTypss: BKL,
ONN, FE GH, ILL not seen, MO, NY, UC, US, VT).
Nuphar advena (Ait.) Ait. . oe ozarkana (Mill. & Standl.) D. Padgett,

nov. BasionyM: Nymph kana Mill. & Standl., Contr. U.S. Natl. Herb.
16:91-92. 1912. Nuphar oza ee 2H (Mill. & Stand. ySeaadh Field Mus. Nat. Hist.
Bot. Ser. 8:310—311. 1931. Nymphozanth ill. & Stand].) Palmer & Steyermark,

Ann. Missouri Bot. Card 22 538. 1935, Re ee ozarkana (Mill. & Stand.)
E.O. Beal, J. Elisha Mitchell Sci. Soc. 72:337. 1956. “ozarkanum’ Type: U.S.A. MISSOURI:
Along the White River in southern Missouri, ms §.2. (HOLOTYPE: US)

As recognized by Padgett (1997), Nuphar pumila sensu lato is a polymor-
phic species within section Nuphar, largely of northern Eurasia, with three
discernible subspecies. This subspecies and the following are restricted to
warm regions in eastern Asia. Nuphar pumila, distinct from the American
N. macrophylla (Padgett 1998), is perhaps the most widely distributed spe-
cies in the genus as a whole

NEES pivade (Timm) DC subsp. oguraensis (Miki) D. Padgett, comb.
ESE v. BASIONYM: Na oguraensis Miki, Bot. Mag. (Tokyo) 48:334—-335.
ae on PE: sr 7, p. 334 (Lecrorype, designated here).

This subspecies refers to a group of relatively rare plants confined to southern
Japan. In the original protologue by Miki (1934) no type was designated.
Although an effort was made to locate original material (four specimens
cited by Miki), specimens of Miki’s collections could not be located in Osaka
(OSA). Therefore, the illustration in the original description (Miki 1934;
Fig. 7, p. 334) of the taxon has been selected to serve as the lectotype. This
plate clearly shows the diagnostic central lacuna of the petiole, a feature
scarcely apparent on herbarium specimens.

Nope pumila subsp. sinensis (Hand.-Mazz.) D. Padgett, comb. et stat.
OV. igs ee Sinense ee -Mazz., Anz. Akad. Wiss. Wien, Math.-Naturwiss.
a 63:8. 1926. Type: CHI HUNAN: icchane sche. in lacunis reg. subtropicae
rsus montem ce s. arenaceo, Handel-Mazzetti 11357 (Tyee: not located,
ce at

ACKNOWLEDGMENTS

My sincere thanks to G. Crow, L. Bogle, T. Lee, D. Les, T. Philbrick, and
J. Sullivan for invaluable advice and assistance; R. Barneby for the Latin

826 Sipa 183)
diagnosis of the new taxon; and J. Wiersema and L. Davenport for critical
review of the manuscript. This work was made possible through research
grants from the International Water Lily Society co the auchor and grants
to G. Crow from the New Hampshire Agricultural Experiment Station and
National Geographic Society. This paper represents Contribution Number
5 of the Ozarks Regional Herbarium.

REFERENCES
Beat, E.O. 1955. Taxonomic revision of the genus Nuphar Sm. Ph.D. Dissertation. State
University of lowa, lowa City, Iowa.
_ 1956. Taxonomic revision of the genus Naphar Sm. of North America
and - urope. J. Elisha Mitchell Sci. Soc. 72:317—346.

1934. On fresh water plants new to Japan. Bot. Gaz. (Tokyo) 48:326—336.
au

ame. = and P.C. Sranpiey. 1912. The North American species of Nymphaea. Contr.
U.S. Nat igre 663-1 08.

Papcert, D. 1 7. A biosystematic monograph of the genus Nuphar Sm. (Nymphaeaceae).
Ph

2D, ae ition, University of New Hampshire, Durham, New Hampshire.

_ 1998. Phenetic distinction between the dwart yellow water-lilies: Nuphar

pimple and N. pumila (Nymphaeaceae). Canad. J. Bot. 76:18.

1. Lesand G.E. Crow. ne — studies of Naphar (Nymp! ),
low water lies: Amer. J. Bot. 83: 35.
,D.H. Les and G.E. Crow is P aes Phylogenetic relationships in Naphar
(ymnsiaecacene): Evidence from morphology, chloroplast DNA, and nuclear ribosomal
DNA. Amer. J. Bc )

Wirrsema, J.H. and C.B. Hettquisr. 1997. Nymphaeaceae. In: Flora North America nort
of Mexico. Vol. 3. Convening ed. N. T. Morin and Editorial Committee. Oxford Uni-
versity Press. New York, NY.

YarskievycuH, G. and J. TURNER. 1990. Catalogue of the Hora of Missouri. Missouri Bortant-

cal Garden. Saint Louis, oe

Y
_

the ye

we

t. 8O.

=)

A NEW HYBRID OF SPIRAEA (ROSACEAE)
FROM OREGON

WILLIAM J. HESS

The Morton Arboretum
Lisle, IL 60532, U.S.A.

whess@mortonarb.org

NICK A. STOYNOFF
Sczence Department, Glenbard East High School
Lombard, IL 60148, U.S.A.

ABSTRACT
A new naturally occurring hybrid between Spiraea douglasii var. menziesii and S. splendens
var. rosea is described and named S. xhitchcockii W. Hess & N. Stoynoff. Morphological,
chromosome number, and pollen stainability evidence is presented. The hybrid is the first
reported triploid for the genus Spraea.

RESUMEN
Se describe un nuevo hibrido natural entre see oe var. menziesii y S. splendens
var. rosea que se nombra S. xhitchcockii W. Hess & N. Stoynoff. Se presentan evidencias
morfolégicas, numero cromosémico y tincién del polen. El une es el primer triploide
citado del género Spiraea.

In the Pacific Northwest, areas where species of Spiraea overlap, hybrids
have been produced (Hitchcock et al. 1961). Hess (1969) presented evi-
dence that S. douglasii Hook. var. menziesii (Hook.) K. Presl and S. betulifolia
Pallas var. /ucida (Greene) C.L. Hitche. hybridized to produce S. x pyramidata
Greene. In at least one area, S. dowglasii var. menziesii and S. splendens (Baumann)
ex K. Koch var. rosea (Gray) Kartesz and Gandhi (syn. S. densiflora Nutt.)
occur together and produce a naturally occurring hybrid herein named Spi-
raea Xhitchcockii.
ape xhitchcocku W. ate & N. Stoynoff (S. ee var. menziesit X

Se var. rosea), hybrida nov. Type: U.S.A. OREGON: Lane Co.: Scott
tee Willamette National Forest, 20 mi W of Three Sc on St. Hwy. 242; mixed
coniferous Gene Abies, Pinus monticola, Tsuga, Juniperus, Salix, Casstope, & Vaccinium;

shrub, scattered around lake, 10-12 dm tall, infl. pyramidal, petals rose-
pink, Elev. 4700', 11 Aug oe Hess & Stoynoff 7283 ae ae MOR; IsoTyPEs:
MO, NA, NY, US)

Differt a Spiraea splendens var. rosea inflorescentia paniculata, pubescentia puberula et Spiraea
douglasit var. menziesii lamina serrulata, calyces lobis puberula, et inflorescentia 1.3—2.8
longiora quam lata.

Sipa 18(3): 827-830. 1999

828 Sipa 18)

—

Shrubs, forming rhizomatous colonies, 10-12 dm tall. Branches pale brown,
becoming dark brown when mature and slightly peeling, lightly puberu-
lent. Leaves (20—)25—45(—60) mm long, (8—)12—18(—21) mm wide; mostly
obovate to narrowly elliptic; lightly puberulent above, puberulent below;
margins serrate, distal 1/2—1/3. Inflorescences pyramidal, 18—75(-95) mm
long, 15—40(45) mm wide; puberulent. F/owers many, in panicles; hypanthia
green, 9-11 mm long, 14-16 mm wide, puberulent within and without;
calyx lobes deltoid, 7-10 mm long, 7-9 mm wide, mostly reflexed, pu-
berulent without, densely puberulent along margins within; petals rose-
pink, broadly elliptic, 13-15 mm long, 11—13 mm wide; stamens many,
pink; carpels 5, separate, follicles glabrous. 7=27.

Additional specimen examined: U.S.A. OREGON. Lane Co.: 21 mi W of Three Sis-
ters off St. Hwy. 242 to Scott Lake, approx. 7 mi W of McKenzie ei 22 Aug 1975, W.
Hess 3628 (MOR)

Distribution.—Scott Lake, in Oregon, is the main locality known to the
authors where Spiraea xhitchcockii occurs. It grows around the lake margin
with its putative parents, S. domglaszi var. menziesii and S. splendens var. rosea.
The putative parents occur elsewhere in the Pacific Northwest and other
hybrids of these species may be found in localities of overlap. Hess (1962)
cited a collection from Pierce Co., WA (Warren 1627, WTU) as a possible
hybrid between the putative parents.

Morphology.—The most obvious morphological difference between Spv-
raea Xhitchockii and its putative parents is a pyramidal inflorescence that is
up to three times as tall as wide, versus the inflorescence of S. dowglasii var.
menziesii that is typically more than three times taller than wide and of
splendens var. rosea, which has a flat-topped or slightly rounded inflorescence

he inflorescences and calyces of 8. xhitchcockii are somewhat puberulent
and not tomentulose as those of 8. dowglasi var. menziesii, or glabrous to lightly
puberulent as that of S. splendens var. rosea. The leaves of 8S. xhitchcockii are

mostly obovate to narrowly elliptical, 2.5—4.5 cm long, 1.2—1.8 cm wide,
finely serrulate, and finely puberulent; those of S. dowglasi1 var. menziesii oblong-
elliptic to obovate-lanceolate, 3—10 cm long, 2—8 cm wide, serrate, and glabrous
to lightly pubescent; and leaves ovate-oval to oblong-elliptic, 2—5 cm long,
1-4 cm wide, serrulate to serrate, and finely puberulent for S. splendens var.
rosea. The intermediate shape of the inflorescence of S. x hitchcockii, when
compared with the putative parents, suggest a hybrid. Spiraea xhitchcockiz
shares the leaf vestiture, length, serrulate margins with those of S. splendens
var. rosea and its leaf shape approaches that of 8. douglasii var. menziesi1. The
shared features of the putative parents suggest the hybrid nature of 8. xAitche gckii.
Cytology.—Hess (1969) reported the chromosome numbers for Spiraea douglast
var. menziesii as n=18 and §. densiflora (= S. splendens var. rosea) as n=9. re

Hess AND StroyNorr, A new hybrid Spiraea 829

A B

b,ef
°
a
e

‘S
o)

Fic. 1. Camera lucida drawings of chromosomes from microsporocytes of Spiraea (X 1000).

A. Spiraea splendens var. rosea, Hess & Stoynoff 7266; a a 1, 94, B. Spiraea douglasii var.

menztesit, Hess & Stoynoff 7281; Metaphase 1, 181. C., D. Spiraea xhitchcockii, Hess & Stoynoff
283. C. Metaphase I, 41,17;. D. Metaphase I, 4y,, 171.

then, all of the chromosome counts of Spiraea in the new world were tetraploids.
The count for S. splendens var. rosea was the first record of a diploid Spiraea
in the new world. The collecting of flower buds, their preservation, and
slide preparation procedures followed Hess (1969). All counts were made
on a Zeiss Photomicroscope II with a camera lucida attachment. Chromo-
some counts were made from microsporocytes of S. splendens var. rosea, S.
douglasti var. menziesti, and S. x hitchcockii. Counts of n=9 for S. splendens var.
rosea confirmed the earlier diploid record (Fig. 1, A). The tetraploid num-
ber of 18 bivalents obtained for 8. douglasii var. menziesii (Fig. 1, B) also

830 Sipa 18(3)

confirmed an earlier count of Hess (1969). Camera lucida illustrations of
chromosomes are shown for S. XAztchcockii (Fig. 1, C, D). The chromosomes
for the hybrid had mostly univalents evident and a few bivalents. The hap-
loid number is 7=27. Spiraea xhitchcockii would appear to be the first known
triploid for the genus.

Pollen stainability —Pollen grains were immersed in cotton blue-lactolphenol
to test for pollen stainability (interpreted as pollen viability) based on pro-
cedures in Hess (1969). The percentage of pollen grains of Spiraea douglas
var. menziesii stained was 84—95 %, similar to Hess’s report for the same species
in 1969. 84-98% of the pollen grains of S. sp/endens var. rosea were stained,
similar to what Hess reported for the same species in his thesis (1962). Spi-
raea Xhitchcockii pollen grains stained from 2—14%. Low pollen stainabil-
ity (i.e. viability) is suggestive of hybridity.

Etymology.—Spiraea x hitchcockii is named in honor of the senior author's
first mentor in taxonomy, C. Leo Hitchcock, one of the best taxonomy teachers
of his era, a student par excellence on the flora of the Pacific Northwest,
and a world class taxonomist.

ACKNOWLEDGMENTS

The authors thank Floyd Swink for his comments concerning this manuscript
and reviewing the Latin description.

REFERENCES

Hess, W.J. 1962. M.S. Thesis. A taxonomic study of Spiraea pyramidata Greene. University
of Washington, Seattle, WA
969. A taxonomic study of Spiraea pyramidata Greene (Rosaceae). Sida
3:298-308.
Hircncock, C.L., A. CRonquist, M. Ownsey, and J.W. THompson. 1961. Vascular plants
of the Pacific Northwest 3. Saxifragaceae to Garryaceae. University of Washington Press,
Seattle, WA

A NEW COMBINATION IN CROIZATIA
(EUPHORBIACEAE)

L.J. DORR
Department of Botany, MRC-166
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560-0166, U.S.A.
dorr!@nmnh.si.edu
ABSTRACT
The unispecific genus Psendosagotia is placed in synonymy under Crozzatia and a new
combination, Croizatia brevipetiolata (Secco) Dorr, is proposed for its sole species. An emended
description is provided for C. brevipetiolata, a dioecious species that was known previously
only from female flowers and fruits. Cro/zatia brevipetiolata is compared to C. neotropica and
C. naiguatensis, two species from coastal Venezuela, and to C. panamensis, a species from
Panama and Pacific Colombia.

RESUMEN

Se ubica el género — Pseudosagotia c indni lel género Croizatia y se

g
propone una nueva combinaci6n, Croizatia brev sais (Secco) ne para su unica especie.
Se presenta una descripcién aa de C. brevipetiolat a, de la cual previamente
s6lo se conocian las flores femeninas y los frutos. Se compara iG. ee con C, neotropica
y C. naiguatensis, dos especies de la region litoral de Venezuela, y con C. panamensis, una

especie de Panama y el extremo occidental de Colombia.

When Secco (1985) described the genus Psexdosagotia, he failed to com-
pare it to Crozzatia Steyerm., which is not surprising given that the latter
genus was then known from two imperfect collections that served as the
types of two species. Had Secco read Steyermark’s (1952, 1978) descrip-
tions of these species or seen the accompanying figures he undoubtedly would
have described his species of Psewdosagotia as a new species of Croizatia be-
cause the two genera are identical in all essential characters save one, which
appears to have been misinterpreted. Subsequently, Webster et al. (1987)
reviewed the genus Crozzatia, which they placed in the Phyllanthoideae,
and they described a third species. Most recently, Webster (1994) placed
Croizatia in the Oldfieldioideae and created a new tribe, Croizatieae, to
accommodate the genus, the correct placement of which he continued to
debate. Webster and collaborators overlooked Psewdosagotia and its sole species
and they cited no other generic synonyms of Croizatia

Salient characters that Secco (1985) observed for Preudieseune: which also
define Croizatia, included the following: pistillate flowers pedicellate; ca-

Sipa 18(3): 831-836. 1999

832 Sipa 18(3)

lyx 5-lobed, petaloid; petals 5, minute, alternisepalous; ovary 3-locular, with
2 ovules per locule; ae -branched, each branch bifurcated; fruit capsular,

with persistent sepal lobes; seeds ecarunculate. In addition, while not men-
tioned by Secco (1985), his illustration (Fig. 1) showed that the sepal lobes of
Pseudosagotia are reflexed in fruit like those of Cro/zatia. Staminate flowers
of Psendosagotia were not available to Secco (1985), but these flowers (de-
scribed below) also agree in shape, size, and ornamentation with those of

—

Crorzatia. Similarly, pollen of Psewdosagotia (described below) agrees with
published descriptions (Webster et al. 1987; Levin & Simpson 1994; Simpson
& Levin 1994) of pollen of C. naiguatensis Steyerm. (The number and nature
of the pores of Pseadosagotia pollen were not visible with light microscopy).
The only character of Pseudosagotia cited by Secco (1985) that does not agree
with Croizeatia (as defined by Webster et al. 1987; Webster & Huft 1988; Webster
1994) is the presence of endosperm, which has not been reconfirmed.

Croizatia Steyerm., Fieldiana, Bot. 28(2):308. 1952. Type: Croizatia neotropica

Steyerm,

ol. Mus. Paraense Emilio Goeldi, Sér. Bot. 2:23. 1985. Type: Psendosagotia
pine Sectors: = Croizatia brevipetiolata (Secco) Dorr.

Croizatia | pe i (Secco) Dorr, comb. nov. BasionyM: Psendosagotia brevipetiolata
Secco, Bol. Mus. Paraense Emilio Goeldi, Sér. Bot. 2:24, nae 1, 2. 1985. Type:

VE aan TRUJILLO: entre Boconé y Guaramacal, 2100-2300 m, 4 Sep 1966

(@ fls., immature fr.), Steyermark & Rabe 97322 (HoLtoryre: NY n.v.; isorype: US!).

Tree, 3—-6(-15) m tall; leaf blades elliptic to lanceolate or oblanceolate,
long-acuminate at apex, Nees at base, (S—)8.5—-21 cm long, (2-)
3—6.5 cm broad, inconspicuously strigose or glabrous beneath, lateral nerves
ca. 8—9 on each side, prominulous with the fainter and more delicate ter-
tiary veins, glabrous above; petioles — mm long, slender, canaliculate,
strigose or glabrous; stipules subulate, ca. 4-5 mm long, scarious, sericeous,
deciduous. Staminate flowers in dense i clusters; pedicel 5-8 mm
long, strigose; sepals 5, elliptic, entire, more or less equal in size, 5.5—6
mm long, 4—5 mm broad, whitish-cream or yellowish-white; petals 5, obovate,
subentire, ca. 1.25—1.8 mm long, ca. 1-1.2 mm broad, glabrous adaxially,
densely sericeous abaxially; disk cupuliform, ca. 1 mm tall, 4-5 mm in diam.,
glabrous; stamens 5, free, filaments ca. 3 mm long, hirsutulous basally; anthers
elliptic, ca. 1.5—2 mm long; pistillode 3—4-lobed, 4—5 mm tall, hirsutu-
lous basally. Pollen spherical, ca. me ktm in diam., and echinate (Cve//o et al.
992, US!). Pistillate flowers in 2—3-flowered axillary clusters; pedicel scri-
gose, |2—28 mm long at anthesis er in fruit; sepals (4) 5, elliptic, slightly
carinate (hooded), 8-10 mm long, 3—5 mm broad, unequal in size (one larger
than the others), strigose adaxially near the base, strigose abaxially, green-
ish, becoming reflexed and persistent in fruit; petals (4) 5, elliptic, ca. 1.25

Dorr, A new combination in Croizatia 833

mm long, ca. 0.75 mm broad, glabrous adaxially, densely sericeous abaxially,
disk cupuliform, ca. 1 mm tall, ca. 5 mm in diam., glabrous; ovary seri-
ceous, hairs white; styles spreading, ca. 3.5 mm eae: twice bifid, the un-
divided portion ca. 1.5 mm long, the primary branches ca. 1.5 mm long,
ultimate tips ca. 0.5 mm long. Capsule oblate, 3-lobed, 1—-1.5 cm tall, 1.5—
1.8 cm in diam., reticulate, strigose to glabrate; columella ca. 7-10 mm
tall, wings persistent. Seeds ovoid, flattened on one side, ca. 10 mm long,
ca. 6—8 mm broad, testa smooth, hilum medial, exalbuminous (Cvel//o et al.
990, US!), cotyledons folded (contortuplicate).

Distribution and ecology.—Locally common on the slopes of the Andes in
Barinas, Lara, Portuguesa, and Trujillo states, Venezuela, where it occurs
from 1400—2300 (—2600) m elevation in lower montane (cloud) forest of-
ten dominated by Wettinia praemorsus (Willd.) Wess. Boer (Palmae). In the
Ramal de Guaramacal (Trujillo state) Cuello (1997) found that Croizatia
brevipetiolata was the most abundant species in vegetation plots she estab-
lished at 2100 and 2300 m elevation. Also, apparently frequent at 1650 m
elevation in dense forest on the slope of Pico Renjifo, Meta, Colombia. In
Venezuela, flowering from (April) May through July and fruiting in Octo-
ber and November. In Colombia, known only from fruiting material col-
lected in June.

In some Venezuelan populations, male plants appear to be less common
(less conspicuous?) than female plants (fide Dorr & Barnett 8046).

Additional material examined. VENEZUELA. Barinas: Dtto. Pedraza, NE of Alto de
La Aguada (ca. 8° 37'N, 70° 40'W) in an area known —— as “Montafias de Tierra Blanca,”
18 Apr 1988 (fl bud), Dorr et al. 4858 (NY, PORT—2 sheets, VEN). Lara: Dtto. Moran, las
cabeceras del Rio Tocuyo, 20—21.5 kms al sur de Humocaro Alto, hacia Guaité, 13 Oct 1974
(fr), eee is eae Espinoza 111112 (VEN), carretera ees Humacaro Alto hacia
Guaité, 14 Nov 4 (fr), van der Werff & Rivero 7915 i. PORT). Portuguesa: Deto.
a. ee Are al Guaramacal, Sector E] Paramito, eee Real La Aguadita—E]
Batatal (ca. 9° 19'N, 70° 04'W), 8 Jul 1b i Dorr e ee 8555 (PORT, US). Trujillo:
Dtto. Boconé, 25 km al SE de Boconé, 26 Jul 1984 ( fils), aes al, 2948 (NY, PORT,
US); Parque Nacional Guaramacal, vertiente norte, 17-18 Jun 5 (2 fls), Cuello et al.
990 (MO, US), Ibid, 17—18 Jun 1995 (6 fls), Cuello et al. oe US); Parque Nacional
Guaramacal, road from Boconé to Guaramacal, SE of Boconé, N slope of mountain, 13 Ju
1995 (@ fls), Dorr & Barnett 8044 (K, NY, PORT, US, VEN), Ibid, 13 Jul 1995 (s fls),
Dorr & Barnett 8046 (DAV, F, PORT, US, VEN); Pdramo de Guaramacal, SE of Boconé on
road from Boconé to Guaramacal, 19 Oct 1990 (fr), Dorr et al. 7437 (NY, PORT); Parque
Nacional Guaramacal, road from Boconé to Guaramacal, SE of Boconé and just above park
Soe. N slope of mountain, 5 Jul 1995 (¢ fls), Dorr et al. 7963 (CANB, K, MO,
Y, PORT, US, VEN); Parque Nacional Guaramacal, en la vertiente norte, 27 May 1995
(9 ass oe ao ee 151 (COL, MO, PORT, US), Ibid, 27 May 1995 (d fils), poe G
Cuello 152 (D PORT, US), Ibid, 15 Jul 1995 (@ fs), Licata & Nitto 304 (COL, PORT,
US). Deto. ate Burbusay, Sector El Alto, 27 Jul 1995 (1), Licata & Gonzdlez 412 pases
COLOMBIA. Meta: Sierra de la Macarena, Central Mountains, Pico Renjifo, 22 Jan 1950
(fr), Philipson et al. 2154 (US)

—

834 Stipa 18(3)

@ C. brevipetiolata

@ C. naiguatensis

@ C. neotropica

* C. panamensis
a ‘s

Fic. 1. Distribution of Cro/zatia species (Euphorbiaceae) in Central and northern South

America.

The paucity and incompleteness of the available material make interpreting
the species boundaries of Cro/zatia difficult. Crotzatia brevipetiolata is very
similar morphologically to C. naiguatensis, which is known from a few col-
lections from the Cerro Naiguata in the Coastal Cordillera of the D.E, Venezuela.
Croizatia brevipetiolata differs from C. naiguatensis, however, in that the leaf

Dorr, A new combination in Croizatia 835

apices are long-acuminate (versus acute or obtuse) and the leaf bases are long-
attenuate (versus attenuate); the staminate flowers are larger in all measur-
able characters (pedicels S—8 versus 3-4 mm long; sepals 5.5—6 x 4-5 ver-
sus 4—4.5 X 2.5—3 mm; disk 4—5 versus ca. 3 mm in diam.; filaments ca. 3
versus 2.2—2.5 mm long; anthers 1.5—2 versus 1.2—1.4 mm long; and pistillodes
4—5 versus 1.8—2.2 mm tall); and the pedicels of pistillate flowers are longer
both in flower and fruit (12—28 versus 8-12 mm long). Collectors’ notes
also indicate that the perianth of pistillate flowers of C. brevipetiolata is greenish
in color (fide Licata & Cuello 152, Licata & Nitto 304, etc.), while that of C.
naiguatensis is cream-colored (fide Berry et al. 4121, 4125, US!). Both C.
brevipetiolata and C. naignatensis can be distinguished from C. panamensis G.L.
Webster by having staminate flowers with free (versus connate) stamens.
Likewise, the two species can be distinguished from the enigmatic C. neotropica
by having shorter (8—28 versus 35—45 mm long) pistillate flower pedicels.
Good flowering material (both staminate and pistillate) of C. neotropica is
still lacking. Croizatia brevipetiolata and C. naiguatensis also can be distin-
guished from the other two described species by having fewer (7—10 versus
12—15) lateral nerves on each side of the midvein of the leaf blade.

The few collections of Cro/zatia begin to suggest distinct geographic ranges
for the species (Fig. 1). Croizatia brevipetiolata evidently is restricted to for-
ested slopes of the northern Andes at elevations ranging from 1400—2300(—
2600) m. Croizatia naiguatensis and C. neotropica are both found in the Coastal
Cordillera of northern Venezuela, but the former occurs in cloud forest at
1900-2150 m in the central portion of this cordillera, while the latter oc-
curs at LOOO—1350 m in the eastern portion of the same cordillera (Steyermark
1978). Croizatia panamensis has been collected only in primary forest at (50—)
300-500 m in Panama and the Chocé region of neighboring Colombia. Additional
material from Amazonian Ecuador and Peru, which Webster et al. (1987) mentioned,
may represent new species and could extend the range of Crozzatia further
south, but none of this material was examined for this note.

ACKNOWLEDGMENTS

lam indebted to Kenneth Wurdack for his observations on the identity
of Psendosagotia (although we part ways on the distinctiveness of Croizatia
brevipetiolata), Basil Stergios for facilitating research in Venezuela, INPARQUES
for granting permission to work in Guaramacal, and the Wildlife Conser-
vation Fund of the National Museum of Natural History (supported by Molson
Breweries U.S.A., Inc.) for funding field work. The curators of NY, PORT,
and VEN were most gracious in lending or permitting me to study speci-
mens in their care. Lynn Gillespie and Geoffrey A. Levin reviewed and made

useful suggestions for improving the manuscript.

836 Sipa 183)
REFERENCES

CurL_o A., N.L. 1997. Floristic diversity and structure of the montane forests of Cruz Carillo
National Park in the Venezuelan Andes. M.S. Thesis, University of Missouri-Sc. Louis,
St. Louis.

Levin, G.A. and M.G. Simpson. 1994. Phylogenetic implications of seo oo
in the Oldfieldicideae (Euphorbiaceae). Ann. Missouri Bot. Gard. 81:2

Sacco, R. de $. 1985.

1985. Pseudosagotia: um novo género de Euphorbiaceae para a ere.
Venezuelana. Bol. Mus. Paraense Emilio Goeldi, Sér. I

ot. 2:23—27.
i ae M.G. and G.A. Levin. 1994. Pollen ultrastructure of the biovulate Euphorbiaceae.
Inc. J. Plane Sci. 155:313—34
ee RMARK, J.A. and COLLABORATORS. 1952. Botanical exploration in Venezuela—II. Fieldiana
Bot. 28(2):243-447.

and )
mountains, Venezuela. Batons 30: = 49,

WessTER, G.L. 1994. ees of the genera and suprageneric taxa of Euphorbiaceae. Ann.
Missouri Bot. Gard. ¢ —144,.

oe : ae OF Gnas | J. STEYERMARK.,
icsacaealanape es

78. New taxa from the Avila and Naiguata

1987. Systematics of Croizatia
id M.J. ae 88 van synopsis of Panamanian Euphorbiaceae.
Ann. Missouri Bor. Gard. 75:1087—

EIGHT NOVELTIES IN ABILDGAARDIA
AND BULBOSTYLIS (CYPERACEAE)
FROM SOUTH AMERICA

ROBERT KRAL
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
MARK T. STRONG
Department of Botany, MRC/166
Smithsonian Institution
Washington, D.C. 20560-0166, U.S.A.
strong. mark@nmnh.si.edu
ABSTRACT

One new species Se A. papillosa, and seven of Bu/bostylis, are described and
illustrated, and their relationships to closely allied species are discussed. In addition,
updated description ie. genus along witha key to the species of New World Abildg andi 1

is provide

RESUMEN

—

Se describe e ilustra una nueva especie de Abi/deaardia, A. papillosa, y siete nuevas especies
de Bulbostylis, y se discute su relaci6n con especi -anas. Ademias, se ofrece una descripcién

actualizada del género, acompafiada de una clave para identificar las especies del Nuevo

Mundo de A bildgaardia.

In our ongoing field-and-herbarium work with American fimbristyloid
Cyperaceae, we have eight new species to report, namely one Abi/dgaardia
and seven Bulbostylis. These novelties are all South American, seven from

the planalto of Brazil, and one from Bolivia. Decisions on placement of species
to genus in this work are based on distinctions between A bi/deaardia, Bulbostylis
and Fimbristylis as given in Kral (1971).

Since the description of North American Abildgaardia given by Kral (1.c.)
has to be amended to include two other taxa, one the new species, the other
a species even today referred to by authors as Fimbristylis bahiensis Steudel,
we give below a modification that now defines the genus in the New World.

Abildgaardia Vahl, Enum. Pl. 2:296. 1805.
Perennial or annual, glabrous, caespitose. Leaves with sheaths closed save

at summit, lacking cilia or fimbriae at orifice, eligulate; blades narrowly

Sipa 18(3): 837-859. 1999

838 SIDA L8(3)

linear to filiform, thickened, flat to variously involute or essentially lack-
ing. Spikelets ovate to linear, either solitary and terminal on wiry culms or
few to several in a loose to dense terminal cluster, rarely in a simple dicha-
sium, in any case subtended by a single involucral bract, this shorter than
inflorescence or spikelet; spikelet scales loosely imbricate, mostly keeled
and subdistichous or arranged so that the spikelet appears compressed and
with torque (twisted), the lower 1—2 scales sterile, those above subtending
perfect flowers or apical ones male, all with dorsal/midcostal zone of 3—5
costae, these convergent at scale apex and either included or excurrent as a
mucro. Florets produced ona short pedicel-joint; perianth absent. Stamens
2—3; filaments flat and twisted; anthers basifixed, narrowly oblong or lin-
ear, the two thecae at maturity longitudinally and laterally dehiscing. Style
3-branched, the base trigonous, smooth or glandular-puberulent, capping
the nipple-like achene apex, dehiscing at achene maturity, upwardly slen-
der, smooth or glandular-puberulent at junction with the linear, glandu-
lar-hairy stigma branches. Achene stipitate-obovoid, the stipe stout, the body
globose to broadly obovoid, strongly tricostate, apically abruptly narrowed
to a pyramidal or truncate-pyramidal nipple, the three convex faces finely
ongitudinally and indistinctly by narrow cancellae, conspicuously

—

lined
roughened by a combination of short, coarse, transverse rugae and/or uni-
form or uneven coarse, large, low and dome-shaped papillae.

There are, by conservative estimate and in the strictest sense, about 15
species of Abildgaardia worldwide in tropical or subtropical regions; only
four seem to be known for the Americas. A key to the American species is
provided below, followed by detailed descriptions of the two which have
bladeless leaves, one of which is new to science.

KEY TO NEW WORLD ABILDGAARDIA

. Plants perennial with well-developed leaf blades; culm bases swollen; spikelets
ovate.
2. Spikelets pale, frequently solitary at apex of culms or 2—3 per culm, in

which case scattered toward scape apex |. A. ovata
2. Spikelets red-brown, mostly few in clusters at apex of culms or rarely solitary
2. A. mexicana
1. Planes wich no leaf blades or at most a cusplike blade no longer than 5 mm;
cum bases not swollen; spikelets linear
3. Scape and leaf sheath surfaces smooth; apices of leaf sheath and spikelet
scales narrowly acute to acuminate; spikelets (S—)17—15 mm long; anthers
1.S—-2 mm long 3. A. baeothryon

3. Scape and leaf surfaces uniformly papillose; apices of leaf sheath and spike-
_

let scales emarginate; spikelets 15-20 mm long; anthers ca. 3 mm long.
4

4. A. papillosa

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 839

1. Abildgaardia ovata (Burm. f.) Kral, Sida 4:77. 1971. Type: from Java, coll.
ign. (HOLOTYPE: G, n.v.). Carex ovata Burm.f., Fl. Indica 194. 1768. Fimbristylis ovata
(Burm. f.) Kern, Blumea 15:126. 1967.

Cyperus es L., Manc. Pl. 180. 1771 oe monostachya (L.) Vahl, Enum.
PL...22296, 1805, Barns ae, - = aia Pl. Jav. Rar. 61. 1848. Iriha
monostachya . ) Kuntze, Rev. Gen. Pl. 751

This is the generic type from India. It is the only cosmopolitan taxon of

the four treated herein. In the Americas, it ranges from peninsular Florida,
U.S.A. southward through the West Indies and Mexico through Central
America to southern South America (Argentina). It is a plant of basic or
calcareous substrates in savannas, grasslands, landward edges of brackish or
salt marsh, calcareous outcrop areas, and on dryish to seasonally dry sites,
mostly at low to medium (1500 m) elevations.
2. Abildgaardia mexicana (Palla) Kral, Sida 4:71. 1971. Type: MEXICO:
Bords de l’Alteseca, prés de Puebla, 13 Jun 1907, FE Arsene s.n. (HOLOTYPE: GZU,
n.v.). Frmbristylis mexicana Palla, Oesterr. Bot. Z. 58:390—391. 1908.

Fimbristylts crassipes Boeck., Linnaea 38:392. 1874, nom. illeg., non Boeck., Flora 41:602
1858.

So far as is presently known, this species is confined to the grasslands of
the Mexican high plateau in Aguascalientes, Durango, Guanajuato, Guerrero,
Distrito Federal, Jalisco, Mexico, Michoacan, Pubela, Queretaro, San Luis
Potosi and Zacatecas. It is found on moist to wet, heavy dark clay earths,
typically at elevations over 2000 meters.

It is noteworthy that the above two taxa have achenes over twice as long
as those of the following two, but are otherwise so similar to them that drawings
of them made equal in size would seem to be of the same species.

3. Abildgaardia baeothryon St. Hil., Voy. Distr. Diam. 2:389. 1833. (Fig.
1). Type: SOUTH AMERICA. Brazi.: “... entre les arbrisseaux de la langue de terra
du lac de Saquaréma, j’indiquerai les suivantes,” Saint-Hilaire s.n. (HOLOTYPE: P, n.v.).

Cyperus geminatus Schrad. ex Roem. & Schult., Mant. 2:95. 1824, nom. oo non aa D
eee ex Ainslie, Mat. Med. Hindoostan 250. 1813. [Abildgaardia g
. & Schult.) Schrad. ex Nees in Mart., Fl. Bras. 2(1):7 1. 1842, nom. J
(Abilene eas Nees, Linnaea 9:289. 1834, nom. nud.] Migs anny
(Nees) L ees in Mart., Fl. Bras. 2(1):71. 1842.
Abildeaardia. ee Kunth, a Pl. 2:248. 1837. Iviha aphylla (t h) Kuntze, Revis.
Gen. Pl. 753. 1891

[Abildeaardia ternata Schrad. ex Nees in Mart., Fl. Bras. 2(1):71. 1842, nom. in syn. |

Fimbristylis bahiensis Steud., Syn. Pl. Shion 2:108. 1855

[Cyperus scirpinus Salzm. ex Steud., Flora 33:229, 1850, nom. nud.; Steud., Syn. Pl. Glumac.
2:108. 1855, nom. in syn.}. (Prana serrpinus Steud., Flora 33:229. 1850, nom.
nud.; Steud., Syn. Pl. Glumac. 2:108. 1855, nom. in syn.].

S40 SIDA 18(3)

Fic. 1. Abildeaardia baeothryon Sc. Hil. (from Kral et al. 72826).—a. Habit sketch.—b. Sec-
tion of culm showing vencral and dorsal view of leaf sheath.—c. Basal sheaths.—d. Inflores-
cence.—e. Dorsal and lateral view of spikelet scales.—f. Style (left) and stamen (right).—g.

Achene.

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 841

Pimbristy lis martit Boeck., Linnaea 37:24. 1871. Iriha martii (Boeck.) Kuntze, Revis. Gen.

Annual or short-lived perennial, densely caespitose, 10—30 cm tall, gla-
brous. Roots slender, fibrous. Leaves few per culm, essentially sheath, dis-
tally opening with scarious entire borders convergent, the dorsal costae forming
an acuminate tip or a cusplike blade no longer than 5 mm. Culms stiffly
erect, slightly twisted, multicostate, smooth, the ribs interrupted by some
deep sulci. Inflorescence terminal, of 1—4(—5) linear, sessile spikelets, 8-15
mm long, subtended by 2-3 short, persistent triangular bracts 3-4 mm
long; spikelet scales arranged subdistichously in a long spiral; fertile scales
ovate-triangular, navicular, 3—4(—5) mm long, narrowly acute to acuminate,
the midzone (keel) thickened, tricostate, pale, the sides stramineous to pale
brown, scarious. Stamens mostly 3, the anthers linear, 1.5—2 mm long, apiculate.
Achene stipitate-obovoid, tricostate, 0.8—1 mm long, faces gray to white,
lustrous, liberally dotted with separate or connected coarse papillae, these
often arranged in interrupted, wavy transverse rugae, the ribs broad but
low, mostly smooth.

Habitat and distribution.—Abildgaardia baeothryon is found on moist to
dryish sands, peats and gravels of savanna and cerrado in the states of Ba-
hia, Distrito Federal, Espfrito Santo, Minas Gerais, Pernambuco, and Rio
de Janiero, Brazil, O-1200 m.

Representative material. BRAZIL. Hab. in siccis prope Ilheos, 1820, Riedel s.n. (US).
Bahia: Sa/zmann s.n. (US); Coastal a 11 km S of Santa Cruz Cabralia, restinga af the
sea, with strand vegetation progressively developing into scrub and low forest further in-
land, 16°22'S, 39°O1'W, sea level, 17 Mar 1974, Harley, Renvoize, Erskine, Bone Braben

and 1 km N along road from the Rio Itanhentinga, restinga, 17°31'S, 39°13'W, sea level,
15 Jan 1977, Harley, Mayo, Storr, Santos & Pinheiro 17972 (US); Serra das Almas, lower NE
slopes of the Pico das Almas, ca. 25 km WNW of the Vila do Rio de Contas, sandstone,
metamorphic and quartzite rock outcrops with associated marsh and damp flushes, 13°33'S,
41°57'W, 1500 m, 17 Feb 1977, Harley, Mayo, Storr, Santos & Pinheiro 19556 (US); Serra
do Sincora, by Rio Cumbuca, ca. 3 km S of Mucugé, near site of small dam on road to
Cascavel, (rae damp sandy soil, sandstone rocks and partly burnt-over vegetation, ca.
13°01'S, 41°21'W, ca. 850 m, 4 Feb ee Harley, meas atiui Brighton & Pinheiro
15947 ee Serra do Sincora, 9 km SW of Mucugé, on road from Cascavel, waste grounc
by Rio Paraguagi, with damp grassland and ae | wo a vd. ca. 13°02'S, 41°25 'W,
950 m, 7 Feb 1974, Harley, Renvoize, Erskine, Brighton & Pinheiro 16101 (US); Serra do
Sincora, S of Andaraf, 16 km along road to Mucugé, near small town of Xique-Xique, sandstone
rocks intersected by small streams, with some disturbed areas by roadside, 12°54'S, 41°19'W,
700-900 m, er Feb 1977, Harley, Mayo, Storr, Santos & Pinheiro 18692 (US); Mun Rio de
Contas, e road to Salto of Rio Brumado, 2 km SE of Rio de Contas, sandy peat of eat in
1. 900 m, 13 Jul 1985, Kral, Wanderley, Cerati & Lima 72737 (SP, VDB);
Mun. Balmieitas, moist sandy intervals in scrub-dotted arenaceous-rocky E-aspect carne
Serra da Larguinha, ca. 15 km § Palmeiras by the Palmeiras-Capao estrada, ca. 950 m, 19
Jul 1985, Kral, Wanderley, Cerati & Lima 72826 (GH, K, NY, SP, US, VDB); In moist sandy

842 Sipa 18(3)

intervals amongst arenaceous boulders and outcrops, campo by Mucujé-Andarai road, ca.
15 km N of Mucujé, just S of Rio Piaba bridge, ca. 940 m, 21 Jul 1985, Kral, Wanderley,
Cerati & Lima 72861 (MICH, MO, NY, SP, US, VDB); Rocky summit, E facing slope just
below TV transmission tower N of BR 242, ca. 5 km W of paved road to Lengéis, damp
sandy sites, ca. 1200 m, 9 Nov 1988, Kra/. Wanderley & Funch 75588 (GH, MO, NY, SP,
TEX, US, VDB); Sandy-rocky campo along Rio Piaba, bridge area, by road to Mucujé from
Andarat, sandy areas between rocks, 900 m, 10 Nov 1988, Kral, i mderley & Lima 75616
(MO, NY, US, VDB, VSC); 5-7 km W of soba 1on road t eira de Freitas, moist
white se es clearings in coastal scrub, 0-50 m, 13 Nov 1988, | W, Wander rley & Lima 75085
(GH , NY, SP, TEX, US, VDB, VPI, we ) ee Federal: Praia de Sernambetiba
ae - Bandeirantes), 23°00'13"S, 43°20'49"W, ao nivel de mar, 4 Apr 1952, Smith

6377 (US). Espirito Santo: Municipio de Vila Vel ha, Lagoa do Milho, beira da lagoa, substrato
ench: pen 14 Jan 1975, Peixoto 369 (US); Mun. Linhares, Re rd. BR-10 )1, Res. Flor. da
Sooretama, solo arenoso imido, 8 Apr 1984, Hatschbach 47722 (US). Minas Gerais: Sandy

areas between sandstone boulders, cerrado ca. 2 km N of Cristalia on road to Grao Mespul:
2 Nov 1988, Kral & Wanderley 75464 (GH, MO, NY, SP, US, VDB, VSC); Campos Rupestres,

BA-Chapada Diamantina, summit near Fumaga Falls, campo rupestre, 16 May 1992, A/ves,
Becker & Roppa 4148 (US). Pernambuco: Prazeres, 7 Sep 1924, Pickel 798 (US); Prazeres,
Licoralgegend, 30 Oct 1932, Pickel 3140 (US); In sandy soil, Recife, Oct 1933, Pickel 3140
(US). Rio de Janiero: 18 Aug 1896, Ule 4155 (US); Municipio de Cabo Frio, Cabo Frio,
Praia do Pontal, 22°56'48"S, 42°01'54"W, ao nivel do mar, 17 Apr 1952, Smith 6563 (US).

4, sae ae papillosa Kral & M. Strong, sp. nov. (Fig. 2). Type: SOUTH
CA. Brazit. Bahia: Mun. Palmeiras, sandy peaty seep areas around arena-
ceous mai ers and outcrops, Serra do Larguinha (Sincord?) ca. 20 km $ of Palmeiras
and due E of Capao by trail, ca. LOOO m, 19 Jul 1985, Kral, Wanderley, Ceratr & Lima
72808 (HOLOTYPE: SP; lsoryPes: GH, K, MICH, MO, NY, TEX, US 3375813, VDB,

$C)

A. baeothryon St. Hil. species Brasiliae atfinis, aqua imprimis differc pableibus alcioribus;

paginis foliorum et scaporum papillosis, apicibus vag 2 longioribus

( mm longis).

Plant annual or short-lived perennial, densely caespitose, 30—40 cm tall.
Roots fibrous. Leaves entirely sheath, the outer ones more scale-like, ovate
to lanciform, to 2 cm long, the principal ones 6-7 cm long, loosely tubu-
lar, papillate, multicostate save on scarious inner band, this distally becoming
the scarious border of the open sheath apex, the margin entire, scabrous,
converging retusely above a short mucro comprised of convergent costae.
Culms slightly twisted, linear, stiff, 0.8—1 mm thick, multicostate, 1-2-
sulcate, the costae fine or coarse, the surfaces dull gray-green, appearing
glaucous because of dense papillosity. Inflorescence terminal, usually of 1
(rarely 2) lance-linear to elliptic-linear spikelet(s), 1.5—2 cm long, of many
scales arranged subdistichously in a long spiral; sterile scales 4—5, narrowly

5

ovate to oblong, 3.5—4 mm long, obtuse and slightly emarginate, the dor-
sal area thickened with 3—5 costae converging to a subapical blunt mucro;
fertile scales mostly oblong to elliptic, 5S—6(—7) mm long, apically narrowly

-y

rounded or emarginate, convex, the dorsal area likewise thickened, with 3

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis

a
ce
ee

ee

rete

nS
MAT

a

—s—

0
es 58)

"4

ee A/T

1. by ome
bal

nikdratih

anit ea

Fic. 2. Abildgaardia papillosa Kral & M. Strong (from the type, Kra/ et al. 72808).—a. Habit
sketch.—b. Section of culm showing dorsal and lateral view of sheath.—c. Inflorescence after
ed.—d. Inflorescence (spikelet).—e. Detail of section of
ikel le (left) and fertile spikelet scale

ICI)

spikelet scales and achenes have sh
culm showing papillose surface.—f. Sterile basal spil
(right).—g. Stamen.—h. Achene and style.

844 Sipa 183)

coarse, usually pale costae, the scarious matrix pale brown to castaneous with
a pale border. Stamens 3, the anthers narrowly linear, apiculate, ca. 3 mm
long. Achene stipitate-obovoid, 1—1.2 mm long, apically obscurely trigonous
but angles low, rounded, the faces convex, transversely irregularly pale-rugose
with large variously interconnected papillae, ridges and papillae connected
by gray, finely vertically striolate intervals.

While Abi/deaardia papillosa is obviously closely related to A. baeothryon,
it is at once distinguished by its taller habit, its papillose (rather than smooth)
foliage and culms, its emarginate or retuse sheath apices and fertile scale
tips (versus acuminate or cuspidate), its longer spikelets which are usually
solitary (rather than clustered) at scape tips, and its longer spikelet scales
and anthers. In the field, the papillate surfaces of leaf and scape make the

—

plant appear glaucous.

BULBOSTYLIS Se Enum. Pl. 2:205. 1837, nom. cons. (neither Bv/bostylis
Steven, 1817, nor DC, 1836, nom. rej.).

Stenophyllus Raf., Neog. 4, 1825, nom. rej.

Oncostylis Mart. ex Nees in Mart., Fl. Bras. 2(1):80. 1842.

These plants make up what must approach 100 species, mostly with wiry
culms and leaves and of acidic, moist to dry, upland habitats in temperate
to tropical climates worldwide. They have been considered by some as a
section of Abildeaardia, by others as a part of Fimbristylis. However, in the
Bulbostylis we have studied, there is but one example {B. truncata (Nees) M.
Strong} in which the leaf sheath apex is not fimbriate. All save one of these
tricarpellate species have a persistent tubercle that is seated above a dis-
tinct suture or atop an achenial “neck.” In the latter instance there is a dra-
matic difference in texture and color between the achenial apex and the style
base or tubercle. One becomes accustomed in the field to note a distinctive
ecology for most Bu/bostylis, nearly all of which are wiry-leaved-and-scaped
physiological xerophytes, denizens of seasonally droughtly, mostly sandy
and acidic sites, such as grassy uplands and fire-controlled open woodlands
or savanna. Since these Bw/bostylis are so uniform in surface characters and
in ecology, we continue to treat such plants as a distince genus. The species

are given in alphabetical order.

Bulbostylis carajana Kral & M. Strong, sp. nov. (Fig. 3). Tyee: SOUTH AMERICA,
BraziL. Para: Serra dos Carajas, 2 km west of AMZA camp N-5S, 6°4'S, 50°8'W, ca.
ce m, scrubby vegetation on ferric rock outcrops, moist low area on outcrops, 13
a aad ‘ling, Condon, Mesquita, Ribeiro & Marinho 5049 (HoLoryPe: [IN PA; tsoTyPEs:
ee VD

Herba annua, dense caespitosa, subglabra, pumila, 9-15 cm alta. Radices delicatula, tenues.

Folia polysticha, graciles, usque ad 5 cm longa; vaginae tubulosae, late dorsaliter viridicarinata,
paucicostatae, glabrae, marginibus latis, scariosis, rufobrunneolis, ad apicem in laminas

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 845

im”

Fic. 3. oe me Kral & M. Strong (from the type, Sperling et al. 5649).—a. Habit
sketch.—b. Leaf a >. Sector of leaf midblade.—d. Leaf sheath and lower blade, adaxial
view.-e. Leaf a a lower blade, oblique view.-f. Spikelet—g. Fertile spikelet scale.—
h. Sector of scape.-I. Anther.—j. Style and stigmas.—k. Achene

846 Sipa 18(3)

abt upte contractis, longifimbriatis; | laminae capillares e I i -analiculatae

dorsaliter valde paucinervosae, apicem versus triquetrae, anguste acutae. Scan] tenues, “deyiter
torti, }—7—costati, 0.3—0.4 mm crassi, unispiculati. Spicae florosae, lanceolato-ovoideae,
6-9 mm ee acutae; squamis fertilibus, cenuibus, apiece Mabie rufo-brunneolis,

noe a mm mee convexis, mediane tricostatis, ad itis. Stamina 3; anthe

I

}

,ca. 2mm longae, apiculatae

,ca. | mm ee eis

inc jae. valde a neee tricostata, aciitneniee a: faciebus valde transverse rugosis, angulatis
umbonatis. Tuberculum globosum, ca. 0.2 mm longum, atrobrunneum.,

Annual, densely caespitose, subglabrous, low, 9-15 cm tall. Roots deli-
cate, slender. Leaves polystichous, few, slender, up to 5 cm long; sheaths
tubular, broadly dorsally green-carinate, few-nerved, ventrally thin, pale
red-brown, abruptly narrowed into leaf blade, there long-fimbriate; blades
capillary, at base canaliculate, thick-margined, dorsally strongly few-nerved,
triangulate towards tip, narrowly acute. Culms filiform, slightly twisted,
5—7-costate, 0.3—0.4 mm thick, unispiculate. Spikelets many-flowered, lance-
ovoid, 6—9 mm long, acute; scales thin, spirally imbricate, the sterile ones
usually 2, shorter than the fertile, acute, the fertile ones oblong, 3-4 mm
long, broadly rounded, scarious save for median costal area, red-brown, apically
ciliate. Stamens 3, the anthers narrowly oblong. ca. 2 mm long, apiculate,
tetrasporangiate. Achene obovoid, ca. | mm long (tubercle included), strongly
trilobed and tricostate, umbilicate, dark brown, the faces strongly trans-
versely rugose, lustrous, the angles umbonate; tubercle round, ca. 0.2 mm
long, dark brown.

—

Habitat and distribution.—Thus far known only from the type collection.

Bulbostylis cavajana is very similar to the widespread savanna species B.
conifera (Kunth) C.B. Clarke, but is definitely annual, the foliage and culms
smooth, the spikelets lance-ovoid (rather than cylindric), the achene more
lustrous, more coarsely rugose, and umbilicate.

Bulbostylis eleocharoides Kral & M. Strong, sp. nov. (Fig. 4). Typr: SOUTH
MERICA. Botivia. Prov. Iturralde. Depto. La Paz: Luisita, 13°05'S, 67°15'W, 180
m.s.n.m., sabana humeda, al W del rio Beni, 24 Feb 1988, Hasse 899 (HOLOTYPE:

LPB; tsoryee: VDB)

Herbe perennis, 60-80 cm alta subglabrata culmis basi sub-bulbosis, cum foliis exterioribus

squamiformibus, glabris. Folia principalia 25—40 cm longa, vaginis proxime conduplicatis,
glabris, ad apicem pallide longifimbriatis; laminis lineari-filiformis, involutis, 0.7—

latis, vaginis 10-plo longiore. Scapi torti et flexuosi, teretes vel subtriquetrii, ca. 1 mm
crassi, glabri. Spiculae solitariae, ovoideae, 8-10 mm longae, squamis numerosis, spiraliter

imbricatis; squamis steriles plures infimis spiculis longiora vel breviora; squamis fertiles
ellipticae vel obovatae, 4.5—5 mm longae, late acutae, convexae. Stamina tria; anther 8 2.5
mm longis. Achaenia obovoidea, ca. 3 mm longa, obscure trigona, paginis ‘avis, Minute
foveolatis.

Perennial rhizomatous herb 60-80 cm tall. Roots fibrous. Culms closely
set along an imbricate-scaly rhizome. Leaves erect or ascending, 25-40 cm

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 847

i)

NY

jam

Fic. 4. Bulbostylis eleoct les Kral & M. Strong (from the type, Hasse 899),.—a. Habit sketch.—
b. Leaf apex, naa side (left), aaa side (right).—c. Sector of leaf blade adaxal side (left),
abaxial side (right).—-d. Leaf base.—e. Spikelet and upper scape.—f. Fertile spikelet scale —g.

Anther (left), achene (right)

848 Stpa 18(3)

long; the outermost appressed chaffy rhizomal scales, grading up culm to
more erect, scale-like basal culm leaves; principal leaves 25—40 cm long,
the sheaths proximally conduplicate with the convex backs striately multinerved,
glabrous, tan, with broad scarious borders pale-fimbriate at convergence with
blade; blades filiform-linear, to 10 times longer than sheaths, 0.7—1 mm
wide, apex subulate, shallowly concave adaxially, the involute margin thickened,
pale, proximally scabrid, otherwise smooth. Culms twisted, flexuous, ter-
ete to subtriquetrous, ca. | mm thick or wide, glabrous. Inflorescence a single
erect to ascending, ellipsoid, acute spikelet 8-10 mm long, of many tightly
spirally imbricate scales with pale firm centers, brown puberulent sides and
broad, dark brown erose and ciliolate borders; sterile scales several, the lowermost

broadly triangular, carinate-keeled, acute or with midcostae excurrent as a
green cusplike blade, this shorter or slightly longer than the spikelet, those
above gradually peconnng longer, grading into the fertile; fertile scales el-
liptic to obovate, 4.5—S mm long, broadly acute, convex, the midcostal zone
thickened, with pale border and a green median. Stamens 3, the anthers
linear, 2.5 mm long, apiculate. Achene ca. 3 mm long, the body obovoid,
2.5 mm long, obscurely trigonous, pale brown with transversely wavy lines
of minute pits on the concave faces, the three angles broadly rounded and
but slightly raised, the achene apex narrowed distally to a distinct neck ca.
0.2 mm long, this capped by a dark, conic tubercle ca. 0.3 mm long.
Paratypes. BOLIVIA. Depto. La Paz, Prov. Iturralde, Luisita, 13°05'S, 67°15'W, 180
m.s.n.m., sabana humeda, W del Rio Beni, pastizal humedo con gramineas bajas en macollos,
28 Feb 1984, Beck G Hasse 10111 (LPB, US). PARAGUAY. In altaplanitie et declivibus
Sierra de aie 907-08, Hassler & Rojas 10040 (MO)

—

Habitat and distribution. Humid grasslands, savannas, Bolivia and Paraguay.

The name of this species alludes to its significant difference from other
Bulbostylis. Its single spikelet bears a strong resemblance to those of many
Eleocharis except for the unusual cuspidate lower sterile bract. Its achenes
are again similar to Eleocharis in that many of the latter are species with

“necked” achenes. The strongest evidence for such a plant to be Bu/bostylis
is, of course, the strongly fimbriate leaf~sheath apex and the well-devel-
oped lamina above it.

a latifolia Kral & M. Strong, sp. nov. (Fig. 5). Tyee: SOUTH AMERICA.
Brazit. Goias: sandy intervals in rocky sandy cerrado on E side BR 040, ca. 4 km N
of Cs ris male ca. 900 m, 6 nes ne Kral, Wanderley & Pereira 75909 (AOLOTYPE:

SP; Isorypes: MO, US Berne DB).

Planta perennis, caespitosa, 70-90 cm alta, basibus culmorum subbulbosis cum foliis
exterioribus squamiformibus. Folia propria plantam 1/5—1/3 aequantia; vaginis oes convexis,
glabris, ad apicem longifimbriatis; laminis vaginis 4—7-plo longiora, 1.5—2.2 mm latis,
planis vel leviter involutis. Scapi anguste lineares, subtriangulares, ca. | mm crassi, glabri.

Inflorescencia diffusa, anguste turbinata. Spiculae in parvis fasciculis [(1—)43—S(—6) per fasciculum]},

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 849

Wy

ay Ve
2 Hh ins:
eal i
eau
tip

t 4
wae weg gael! “a g
CONC ECL aa
SOUR ME TAPS
A) LeaaPat t
Xi ie ile My)
See aed ee
ey, otha aut Ay joe
Se

3

= 32:
ine
SS
=

Fala i
an ui i

Fic. 5. Bulbostylis latifolia Kral & M. Strong (from the type, Kra/ et al. 75909).—a. Habit
sketch.—b. Leaf apex.—c. Sectors of leaf midblade, adaxial oe abaxial (right).—d. Leaf base,
abaxial view.—e. Young offshoot a plant bas nlarged leaf blade sectors, adaxial (left),
abaxial (right).—g. Inflorescence.—h. oie Fertile snore scale._j. Anther on fila-
ment

nt apex.—k. Style and ae ee, Achene

850 Sipa 18(3)

ideae vel HAASE GSE i~5 mm longae, acutae, brunneae vel pallide ferrugineae.

—

Sones files anguste ovatae, 4-3 mm longae, acuminatae, naviculares. Staminae duo
vel tria; antheris 2 mm longis, en ulatis. Achaenia trigono-obovoidea, trilobata, ca.
1 mm longa (tuberculo includens), subtiliter foveolata; tuberculo conico,

Perennial, caespitose, 70-90 cm tall. Roots diffuse-fibrous. Culm bases
sub-bulbous, firm, with short, scaly bulbous offshoots. Leaves sub-basal,
few per culm, the lowest scale-like, bladeless or with blades shorter than
sheaths; principal leaves 1/5S—1/3 as long as the plant, erect or ascending,
15—25 cm long; sheaths abaxially convex, strongly multi-ribbed, glabrous
with broad scarious red-brown to brown borders acutely converging to blade,
there long-fimbriate; blades linear, 4—7 times longer than sheaths, level or
slightly concave adaxially, 1.5—2.2 mm wide, obliquely broadly acute, greenish-
brown, margins slightly thickened, pale, densely ciliate or ciliolate, sur-
faces smooth save for scabrellous adaxial apex, abaxially multicostate. Culms
narrowly linear, multi-costate, subtriangular, green, glabrous. Inflorescence
a narrowly turbinate compound of variously peduncled fascicles, this much
exceeding short-bladed lanceolate involucral bracts 1-3 cm long; branches
several, mostly ascending around subsessile central fascicles, the longer ones
5-10 cm long, either terminating in single spikelet clusters or there re-
branching in compact or open fashion; spikelets (1—)3—5(—6) per fascicle,
narrowly ovoid or lanceoloid, 4 ae mm long, acute, brown to pale red-brown;
fertile scales 3—6, narrowly ovate, 4-5 mm long, navicular, acuminate. Sta-
mens (2—)3; anthers oblong, ca. 2 mm by long-apiculate. Achenes trigonous-
obovoid, ca. | mm long (tubercle included), apically trilobed, the faces dark-
foveolate-puncticulate, gray, the angles paler, rounded; tubercle conic.

Paratypes. Brazil. Goias: sandy intervals in rocky sandy cerrado on E side BR 040, ca.
4 km N of Cristalina, ca. 900 m., 6 Dec 1988, Wanderley et al. 1877 (SP, VDB); rocky
campo, S—6 km by road N of Alto Paraiso, sandy peat of E and NE aspect, ca. LOOO m, 30
Nov | O88. Kral, Wanderley, Cavalcanti & Pereira 75757 (SP, US, VDB); 4 km N of Cristalina
oft E side BR 050 sandy intervals in rocky cerrado, ca. 10 00 m, 6 Dec 1988, Kra/, Wanderley
& Pereira 75909 (SP, US, VDB).

Habitat and distribution —Rocky, sandy cerrado, 900-1000 m, Goias.

This species is distinctly allied to Bu/bostylis junciformis (Kunth) C.B. Clarke
but is readily distinguished by its broader, smoother, ciliate-margined leaves
and fertile spikelet scales with apex not excurved and midcosta not excur-
rent. The grayish achene surfaces are distinctly foveolate-puncticulate, the
pits short-rectangular.

say ee ee aaa Kral & M. Strong, sp. nov. (Fig. 6). Type: SOUTH
AM 3naAzIL. Minas Gerais: Serra do Cipo, Santana do Riacho, afloramento
ie a ena eral proximo a portaria do IBAMA Alto Palacio,
16 Nov 1995, J.A. Lombardi 1029 (HoLoryPE: BHCB; tsorype: VDB).

Planta perennis, 15—30 cm alta, rhizomatosa, rhizomate crasso, caudiciformis, dense piloso,

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis

Sy
Note
y

\

: erseeraesr ty mioer ncnwaesh 3:
eh Ek Et ST DC
ENTREE

axes

eeerener
os
Apo ares MAES
er eres

ahs SeeO RAT ho wot
tore ety te eu ae

Fic. 6. Bulbostylis lombardii Kral & M. Strong (from the type, Lombardi 1029).—a. Habit
sketch.—b. leaf apex.—c. Section through midblade showing adaxial and abaxial surface.—d.
Detail of sheath apices.—e. Inflorescence.—f. Involucral bracts.—g. Fertile spikelet scales and
achene.—h. Achene.

852 Sipa 18(3)

squamoso et fibroso. Folia exteriora Sqeeaeena, margine dense albopil 1OSIS; folia principalia

vulgo expansa, S—8 cm longa; vaginis convexis, prominente 3-5-costatis, ad apicem longe
ciliatis; laminis vaginis 4—6-plo longioribus, rigidis, 0.5—1 mm latis, dorsaliter scabridis.
Scapi erecti, rigidi, subteretes, ca. | mm crassi, dense pallide puberuli. Inflorescentia capitata,

involucrata, late ovoidea vel hemispherica aut subglobosa, 1—1.5 x 1—1.5 cm; bracteae involucralis
extimae ovatae, 5 mm longae, piloso-ciliatae, mucronatae, mucrone 2—3 mm longae; bracteae
involucralis interiores ovatae, 5 mm longae, mucronulatae. Squamae fertiles naviculares,
ca. } mm longae, obtusae vel emarginatae. Stamina tria; antheris ca. 4mm longis. Achaenia
lata obovoidea, triloba et tricostata, 1.2—1.5 mm longa, umbilicata, valde transverse rug-
osa, tuberculo conico, ca. 0.3 mm longo.

Perennial densely caespitose rhizomatous herb, 15—30 cm tall. Roots thickened
(for a Bulbostylis), fibrous. Rhizome stout, knotty, ascending, caudiciform,
the newer surfaces (internodal) densely white-piliferous. Shoots numerous,
densely spirally arranged around caudex, there persisting as an often burnt
stubble of fibers, hairs, and old leaf sheaths, the newer (distal) caudex pro-
ducing rosettes of a season. Outer rosette leaves invested in tufts of white
trichomes, mostly scale-like, mostly sheath, white-pilose-ciliate, thence grading
progressively longer distally on culm, the costae of the dorsal median area
converging to form increasingly longer blades, the scarious sheath borders
with ciliae concentrated more at apex; principal leaves spreading, 5-8 cm
ong, sheaths convex with 3(—5) broad costae and broad, scarious, brown,
apically white-pilose borders; blades 4—6 times longer than sheaths, linear-

—

filiform, 0.5—1 mm wide, adaxially shallowly involute or plane, with pale
incrassate borders, abaxially convex 3(-5)-costate, sparingly scabrid, pale
green. Culms stiffly erect, linear, subterete, ca. | mm thick, several-cos-
tate, costae densely pale-puberulent. Spikelets in dense broadly ovoid to
hemispheric or subglobose involucrate heads 1—1.5 x 1—1.5 cm; involucral
bracts in series, mostly broadly ovate, ca. 5 mm long, brown, ciliate, pu-
berulenc, convex, the lowermost with costal area excurrent as cusp 2-3 mm
long, those above grading shorter to mucros, thence inward to fertile scales,
these oblong to elliptic, navicular, ca. 3 mm long, obtuse or emarginate,
the tricostate scabrellous keel not excurrent, the margin entire. Stamens 3;
anthers linear, ca. 4 mm long, apiculate. Achene trigonous-obovoid, 1.2—
1.5 mm long, apically strongly lobed, umbilicate, each lobe with a pale
crest, the shallowly convex lobe faces dark brown to nearly nigrescent, strongly
transversely rugose; tubercle nearly black, depressed-conic, 0.3 mm long.

Habitat and distribution —This species of the planalto of Brazil is yet known
only from the type.

sa Ago Liao Kral & M. Strong, sp. nov. (Fig. 7). Tver: SOUTH
AN

. BraziL. Minas Gerais: Cerro do Cabral, Armazin de Lag

ane

‘
ae > anc NNW of Joaquim Felicio, ca. 1100 m, sandy paereaene transition,
30 Oct 1988, Kral, Wanderley & Lima 75381 (HoLoryePe: SP; isoryPes: GH, MO, NY,
Us, VibB).

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 853

Fic. 7. Bulbostylis pachypoda Kral & M. Strong (from the type, Kra/ et al. 75381).-a Habit
sketch—b. Leaf apex—c. Sector of leaf midblade, adaxial (left), abaxial (right).—-d. Sector of
leaf midblade (Kral et al. 75175).-e. Leaf base.—f. Upper portion of leaf sheath and scape
base (upper left); idealized sketch of a young outer leaf transitional to inner principal leaf
(lower right).—g. Inflorescence.—h. Lower fertile spikelet scale.—i. Fertile spikelet scale.—j.
Two achenes.—k. Anther.

854 Sipa 18)

Plantae perenne, dense caespitosa, 20—35 cm altae, culmis basi bulbosis, arcte interconnectis,
fola exteriora squamiforma, triangulata, valde nervosa, enn Folia principalia
(S—)8—15 cm longa; vaginis pallide brunneolis convexis, ad apicem parce us fimbriatis
aut glabris; laminis filiformis involutis, vaginis 1—2-plo oe ca. 0.2—0.4 mm latis,
margine remote scabriusculis. Scapi filiforme, a 0.4—0.5 mm crassi, paucicostati,
glabre. Inflorescentia involucrata, pseudocapitata, terminales; fasciculi spicularum 1—2 (aut
inferiore unam spiculam reducti), approximati, ovoidei, turbunati vel himispherici, usque

ad 1.5 mm lati, bracteis infimis setaceis inflorescentiam leviter vel multo excedentes. Spiculae
1 he

ovoideae, 4—5 mm longae, vulgo acutes imbricatis, infimis valde carinatis,

Se acs vel mucronatis; squamae fertiles latae ovatae, 3.5—4.5 mm longae, convexae, costis

3, pallidis, lateribus atrobrunneolis vel castaneis, nitidis, minute puberulentis. Stamina
tria; antheris oblongo-linearis, ca. 2 mm longis, apiculatis. Achaenia obovoidea vel late
obovoidea, ca 1—1.2 mm longa (tuberculo includens), trilobata, tricostata, superficiebus
margaratacea aut cano-eburneis, subciliter longitudine striolatis, costis prominentibus niveis.

Plants perennial, densely cespitose, 20-35 cm tall, the culm bases bul-
bous, closely interconnected at base, forming short thick lines radiating from
clump center. Outer leaves scale-like, triangular, strongly nerved, the broad,
scarious borders long-fimbriate. Principal leaves ascending to spreading,
(S—)8—15 cm long, the sheaths pale brown, convex, sparsely ee or

smooth at the acute to acuminate oe blades filiform, involute, 1-2 times
longer than the sheaths, ca 0.2—0.4 mm wide, abaxially tricostate, - lat-
eral costae forming smooth or remotely scabridulous margins. Culms filiform,
flexuous, 0.4—0.5 mm thick, few-costate, glabrous. Inflorescence terminal
spikelets in an involucrate, head-like, ovoid to turbinate or hemispherical
fascicle, or this directly subtended by an additional but smaller fascicle, sometimes
reduced toa single spikelet. Involucral bracts several, setaceous-tipped, the
lowest bract slightly to much exceeding the inflorescence, those at higher
levels progressively shorter-bladed, those ee individual spikelets
mostly cuspidate or mucronate. Spikelets ovoid, 4—5 mm long, mostly ac acute,
oa several loosely spirally imbricate scales; fertile scales mostly ovate, 3.5—

4.5 mm long, strongly convex, strongly curvate-keeled, the sides deep ted-
brown to castaneous, the keel area tricostate and conspicuously paler. Sta-
mens 3; anthers oblong-linear, ca. 2 mm long, apiculate. Achenes narrowly
to broadly trigonous-obovoid, ca. 1—1.2 mm long (tubercule included), slightly
trilobed, tricostate, the surfaces a lustrous gray-white or pearl, finely longi-
tudinally striolate, the costae strong, smooth, contrastingly pure white.

Paratypes: BRAZIL. Maro Grosso: 5 km E of Primavera by BR 070, sandy-gravelly
campo-cerrado transition, 6 Oct 1988, Kral, Wanderley & Lima 75175 (NY, SP, US, VDB).

The affinities of Bu/bostylis pachypoda are perhaps most with the ubiqui-
tous B. juncoides (Vahl) Kiik. ex Osten, particularly examples from south-
ern Brazil, Paraguay or Argentina, in which spikelets are often dark and
crowded into head-like terminal involucral fascicles. However, B. pachypoda
has t

—

uicker, more bulbous culm bases, these connected in ascending lines

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 855

from a clump center; its leaves and culms tend to be narrower, more smooth;
its achenes, which are paler, are usually a lustrous “pearly” white, and are so
finely striolate as to appear smooth. The young lower (outer) leaves, usually
concealed by a stubble of burned back or dried older foliage, tend to have
completely fimbriate margins and appear to be transitional to the even hairier-
sheathed B. jacobinae (Steud.) Lindm. and allies. The anthers reach 2 mm in
length, which puts them in a range longer than that for B. juwncoides (Vahl)
Kiik. ex Osten and allies.

Bulbostylis scirpoides Kral & M. Strong, sp. nov. (Fig. 8). Tyee: SoutH AMERICA.
Brazil. Gotas: ca. 5 km N of jct. road W to Pires do Rio by BR 050, seep meadow
and associated cerrado by pond, ca. 800 m, seep by pond, 900-1000 m, 7 Dec 1988
Kral & Wanderley 75942 (HoLoryPe: SP; isorypes: US, VDB).

Planta perennis, 60-80 cm alta, c aespitosa, culmis basi bulbosis, caudiciformibus, Se ae

cum foliis exterioribus cquaniioen bre Folia principalia propria plantam 1/3—2/3 aequantia;
laminis filiformis involutis, vaginis 4—6-plo longiora, tricostatis, sage subulatis. Scapi
lineares, subteretes, 0.6—0.7 mm crassi, glabri. Inflorescentia involucrata, receptaculo piloso,

spiculis sessilis, (1—)2—3(—5), ovoideis vel eeeclsidcis: ae squamis fertiles
anguste ovatis vel late ellipticis, curvato-carinoalis, obtusis, ae apicem ciliolatis. Stamina
tria; antheris ca. 3 mm longis. Achaenia lato-obovoidea, ca. 1.2—1.4 mm longa, prominente
tricostata,

Perennial, caespitose, 60-80 cm tall, the shoots arising from bulbous short
stems; receptacle and lower shoot internodes with dense, pale, red-brown
pilosity, 1-2 cm long. Roots fibrous. Lowest leaves scale-like, ovate to lan-
ceolate, to 2 cm long, the inner ones cuspidate with sheath borders pilose
apically; principal leaves ascending to erect, 40—60 cm long, the sheaths to
10 cm long, narrowly convex with 3—5 strong medial costae, the sides with
several fainter costae and scarious red-mottled borders, gradually tapering,
then abruptly joining blade, there long-villous-fimbriate; blades 4—6 times
longer than sheaths, filiform, ca. 0.5 mm thick, adaxially concave with 2
pale costae marginally, abaxially strongly convex with | strong costa, the
marginal costae with sparse, appressed, inward or retrorsely directed stiff
hairs, distally tapering to triquetrous subulate tips. Culms erect, subterete,
ca. 0.6—0.7 mm thick, coarsely 4-6-costate, glabrous. Inflorescence (1—)2—
3(—5)-spicate, spikelets sessile above an involucre, the receptacular surface
sparsely pilose, the lowest involucral bract longest, ovate to lanceolate, tricostate-
keeled, slightly longer to slightly shorter than subtending spikelet(s), slightly
to much exceeding the inflorescence, to 3 cm long, the base ovate, keeled,
long-ciliate, the midcostae excurrent as a mucro, cusp, or linear blade; spikelets
mostly ovate, 8-IO0 mm long, acute, with fertile scales narrowly ovate to
broadly elliptic, mostly 5 mm long, curvate-keeled, obtuse or emarginate,
apically ciliate. Stamens 3, the anthers linear, ca. 3 mm long, apiculate. Achene
body obovoid-trigonous, 1.2—1.4 mm long, rib-angled, pale brown, angles

856 Stipa 18(3)

imm

Fic. 8. Bulbostylis scirpoides Kral & M. Strong (from the type, Kral & Wanderley 75942).—a
Habit sketch.—b. Leaf apex. —c. Sector of leaf at midblade, adaxial view.—d. Leaf sheath, abide

view.—e. Inflorescence.—f. Fertile spikelet scale.—g. Anther.—h. Two achenes.

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 857

smooth, faces shallowly convex, lustrous, shallowly transversely rugulose
with very fine vertical striolae; tubercle either deciduous or short-persis-
tent, oblong, 0.3-0.4 mm long.

Paratyres. BRASIL. Distriro Feperar: Chapada da Contagem, 23 km NNW do Centro
de Brasilia, brejo estacional, a beira da estrada, 19 Apr Age ins 16 (NY, VDB).
Goias: ca. 5 km N of jet. ae W to Pires do Rio by BR 050 meadow and associated
cerrado by pond, ca. 800 m, seep by pond, 900-1000 m, 7 Dec 1988, Wanderley 1910 (SP,
VDB).
Habitat and distribution —Sandy marshy grass-sedge campo, Goias.

his is again an example of that complex of Bu/bosty/is in which the cen-
tral axis of the plant is compact, pilose-vestite, a feature also displayed on
the inflorescence axis (receptacle).

ee spectabilis Kral & M. Strong, sp. nov. (Fi ig. 9). Tyr: SOUTH
RICA. Brazit. Parana: Vila Vellia, abundant in occasional wet sand, -fac-
ing ae covered by savanna type vegetation, 3 Mar 1970, Koyama, Koyama, Hatschbach

& E. de Lima 13848 (uoLorype: NY; isoryre: VDB).

Planta perennis, 4-55 cm alta, caespitosa, subglabraca, crassirhizomatosa, culmis basi bulbosis,

cum foliis exterioribus Seninifenca bine: glabris. Folia principalia 15— cm sae, vaginis
pall ig brunneolis, convexis; laminis filiformis involutis, multi- vel ‘ inis longiore,

ca. 0.3-0.5 mm ais: Scapi lineares, subteretes, ca. 1 mm crassi. iyi ee capitata,
involucrata, ovoidea vel nemisphastich, multispiculata, usque ad 1.5 cm lata, bracteis 1—3
e caudati inflorescentiam excedens; spiculae lanceoloideae, 8-10 mm longae, squamis

jon

vald

fertiles anguste ovatis, 4.5—5.5 mm inne acuminatis. Stamina tria; antherae ca. | mm

longis. Achaenia obovoidea, |—1.2 mm longa, tricostata, minute cancellata.

Caespitose perennial 4—55 cm tall, producing rhizomes which are inter-
connected bulbous bases of culms. Roots fibrous. Outer culm leaves scale-
like, 4-10 mm long, strongly nerved, brown, glabrous; inner leaves pro-
gressively elongating; principal leaves erect to strongly ascending, 15—25
cm long; sheaths pale brown, scarious, multicostate, convex-backed, the scarious,
pale red-brown, friable borders acutely converging to blade, there pale-fimbriate;
blades several to many times longer than sheaths, filiform, 0.3—0.5 mm wide
at midblade, apex filiform-setaceous, triquetrous, subulate, surface pale yellow-
green, essentially glab1 ous save for a few scabrellae toward blade base, adaxially
concave to canaliculate, abaxially mostly tricostate, narrowly sulcate. Culms
linear, several-costate, terete, ca. | mm thick, pale yellow-green. Inflorescence
capitate, involucrate, broadly ovoid to hemispheric, 1—i.5 cm broad; outer
involucral bracts lanciform or narrowly ovate, 5—7 mm long, slightly keeled,

costae excurrent as cusp or narrowed acuminately to filiform-setaceous blades
to 8(—9) cm long; fertile scales narrowly ovate or oblong, 4.5—5.5 mm long,
acuminate, convex-based, apically keeled, with 3 costae. Stamens 3; anthers
elliptic-linear, ca. 1 mm long, apiculate. Achenes narrowly obovoid, trigonous,

858 Sipa 18(3)

Ilmm

\
I
eo

Fic. 9. Bulbostylis spectabilis Kral & M. Strong (from the type, Koyama et al. 13848).—a. Habit
sketch.—b. Leaf blade apex.—c. Adaxial (left) and abaxial (right) sectors of leaf midblade.—
_ Leaf sheath, abaxial side.—e. Inflorescence.-f. Spikelet.—g. Involucral bract.—h. Fertile
i. Stamen.—j. Achene, abaxial faces (above), idealized enlargement of some

ic
spikelet scale.—1. S
cancellae (left), cross-section (right).

KRAL AND STRONG, Novelties in Abildgaardia and Bulbostylis 859

1—1.2 mm long (tubercle included), the adaxial side broadest, abaxial faces
more convex, pale brown, angles dark brown, surfaces minutely cancellate.

Habitat and distribution. —Thus far known only from the type collection

This plant, superficially resembling robust examples of Bu/bostylis sphaerocephala
(Boeck) C.B. Clarke or capitate forms of B. junciformis (Kunth) C.B. Clarke,
differs most significantly in the peculiar rhizome of interconnected bulbous
culm bases, in having perhaps the most glabrous foliage of its complex with
hairs confined essentially to the pilosity of leaf sheath apex, and in the dis-
tinctive involucre, some bracts with setaceous tips to nearly 1 dm long.

ACKNOWLEDGMENTS

We would like to thank Dr. Richard Carter (VSC) and Dr. Dan Nicolson
(US) for reviewing the manuscript; curators at MO and NY for loan and/or
exchange material; José Cedefio-Maldonado for translating the abstract from
English to Spanish; and the National Geographic Society for its travel grant
to R. Kral during 1988. These aids are gratefully acknowledged.

REFERENCES

Kra_, R. 1971. A treatment Cee Bulbostylis, and Fimbristylis (Cyperaceae) for
North America. Sida 4:57—

860 Stipa 18(3)
BOOKS RECEIVED

Scorr, RicHarp W. 1995. The Alpine Flora of the Rocky Mountains.
Volume I. The Middle Rockies. ISBN 0-87480-482-5, hbk). Uni-
versity of Utah Press, Salt Lake City, UT. $110.00. 901 Pp. Line drawings.
Maps.

From the Preface: The flora serves a dual purpose according to the author. Firsc, it deals
with che alpine plants within the political boundaries of the state of Wyoming; second, it
treats the more natural physiographic unit, the Middle Rocky Mountains, which encom-
passes the states of Montana, Wyoming, and Utah, and a very small bit of Idaho,

From the Flora: All the flowering plants and vascular cryptogams known to occur above
timberline in the Middle Rocky Mountains are described in the manual, as are the timber-
line conifers. The flora consists of 700 taxa including 2 supeperl and 314 varieties. The
taxa included in this manual are based on specimens c cced throughout the Middle Rocky
Mountain region. The taxonomic treatment begins with a dichotomous key to all the families.
The families are then presented in alphabetical order for ease of location, and each family
is also organized alphabetically by genus and species. Dichotomous keys are ay ed for
each taxonomic rank, ultimately leading to the species treatment. Infraspecific taxa are

treated within the species. Each species description is based on personal observations of
plants in the field, herbarium specimens, and references from regional manuals and floras.
The species description is followed by a short habitat description. Next the Middle Rocky
Mountain distribution is given by mountain range. Comments on infraspecific taxa and
how to distinguish them follow the distribution information, The distribution maps show

jon

the distribution pattern of each species in the alpine flora. The dots are based on herbarium

specimens and, in a very few cases, field observations when no specimens were available.
Contents include:

Preface

Acknowledgments

Introduction

The Alpine Zone

The Middle Rocky Mountains

The Flora

The Middle Rocky Mountain Alpine Flora

Appendixes, Bibliography

Index to Common Names

Index to Latin Names

Sipa 18(3): 860. 1999

THE FLORA OF INDIAN SHELL RINGS
FROM COASTAL SOUTH CAROLINA
TO NORTHERN FLORIDA

RICHARD STALTER and MARITESS LEYVA
Department of Biological Sciences
St. John’s University
Jamaica, NY 11439, U.S.A.

DWIGHT T. KINCAID
Department of Biological Sciences
Lehman College CUNY
Bronx, NY 10468, U.S.A.

ABSTRACT

Native American shell ae are composed almost exclusively of shellfish remains, and
occupy coastal sites. The nine shell rings and one shell mound selected in this study range

ye ae 40 ere north of Charleston, South Carolina to 10 kilometers north

St. Augustine, Florida. The vascular flora consists of 241 species within 182 genera in

. families. The Poaceae and Asteraceae are the largest families in the flora. Soil salinity

and daily flooding affect the distribution of vascular plants at the tidal shell rings. Calcar-
eous soils influence species distribution at inland shell ring sites. Each taxonomic entry in
the checklist is accompanied by an annotation, which includes frequency of occurrence,

—_—

rarity status and pertinent synonym
Key worbs: Indian Shell Rings, flora, distribution, biodiversity.
RESUMEN

Los “shell rings” nativos de América estén compuestos casi exclusivamente de restos de
ings” y un “shell mound” selec-

—

conchas marinas y ocupan lugares costeros. Los nueve “shell r
cionados en este estudio van aproximadamente desde 40 kilémetros al norte de eee
Carolina del Sur hasta 10 kil6metros al norte de St. Augustine, Florida. La flora vascular cor
siste en 241 especies agrupadas en 182 géneros de 80 familias. Las Poaceae y Asteraceae son
las mayores familias de la flora. La salinidad del suelo y las eae igi afecta cae a distri-
bucidn de las plantas vasculares en los “shell rings” intermareales

la distribucién de las especies en los “shell ring” interiores. Cada entrada taxonémica del catélogo
se acompania de una anotacion de la frecuencia, estado de rareza y los sinénimos pertinentes.

INTRODUCTION
The shell rings of the South Atlantic Coast are ring or arch-shaped ridges
consisting primarily of shellfish remains, which were constructed by pre-
historic Native Americans. They are located approximately between 32°
55' N (ewee Shell Ring, SC) to 30° 05' N (Guana River State Park, FL).

Stipa 18(3): 861-875. 1999

862 Stipa 18(3)

All known shell rings are located on estuaries or tidal creeks. They occupy
high ground immediately adjoining salt marshes or occasionally are iso-
laced in salt marshes a few hundred feet offshore. Interiors of shell rings at
tidal sites are marshy, while the higher sites usually support a heavy growth
of vegetation. The circular shell ridges range from about 33 to 66 m in outside
diameter, 0.5 to 2.5 m in maximum height and 6.4 to 17.8 m in basal width.
They are covered with vegetation and are composed almost exclusively of
shells, mainly those of oysters. Shells of periwinkle, knobbed whelks, clams
and mussels are present in reduced amounts (Dorroh 1971; DePratter 1976;
Trinkley 1985).

Rings occur in complexes as well as in isolation. The largest known ring
is at Sapelo Island, Georgia and is associated with two smaller rings. At the
Skull Creek Shell Ring, Hilton Head Island, South Carolina, the rims of
two rings are superimposed at one point (Hemmings 1970). Shell speci-
mens from rings in South Carolina and Georgia have been radiocarbon dated
and are approximately 3900 to 3100 years old (Calmes 1967; Waring &
Larson 1968

Numerous uses for the rings have been postulated, including ceremo-
nial, recreational, and exploitative functions, e.g. fish traps, (Edwards 1965).
Recent investigations suggest that at least some of the rings were habita-
tion sites, with the rings gradually developing from kitchen refuse associ-
ated with house middens arranged in a circular fashion (DePratter 1976;
Trinkley 1985). The relatively clear interiors of the shell rings may have
functioned as areas of communal activity (Trinkley 1985).

The objective of this study is to determine the vascular flora at shell rings
on the coast of South Carolina, Georgia, and Florida, the only sites on the
east coast where shell rings are known to exist. Several calcareous plants,
including the rare Sageretia minutiflora, grow almost exclusively at these sites.

Several well known individuals have examined shell mounds or rings along
the Atlantic coast. William Bartram described the flora of shell mounds
along the coast of South Carolina bordering the sounds and inlets near Hilton
Head in 1773. Among the plants listed by Bartram at shell mounds were
Magnolia grandiflora, Pinus taeda, Laurus borbonia (Persea borbonia), Quercus
sempervirens (Quercus virginiana), Corypha palma Sabal palmetto), Prunus laurocerasus,

Llex aquifolium (1. opaca), and Juniperus americana (Juniperus virginiana).
There are several studies of the vascular flora of shell mounds along the
southeastern and Gulf coasts of the United States. Brown (1936) reported
that the most common plants found on Louisiana shell mounds are Juncus
roemerianus, Spartina patens, Scirpus olneyt, Fimbristy lis castanea, Iva frutescens,
and Salicornia spp., all salt marsh species. Laessle (1942) noted that certain
plants occupying the Orange Point shell mound in Florida, “seem to per-
sist on the highly calcareous soil,” and are found, “nowhere else in the area.”

STALTER ET AL., Flora of Indian shell rings 863

Griffen (1948) reported that the Florida shell middens are “extremely in-
teresting,” and that they support, “a more or less specialized vegetation.”
Kurz and Wagner (1957) studied the distribution of salt marsh vegetation
along an elevation gradient at the Buzzards Island shell ring, South Caro-
lina, but did not recognize the site as a shell ring. Eleuterius and Otvos
(1979) surveyed a shell mound at Cedar Island, Mississippi, and reported a
vascular flora of 62 species. They reported that a el of plants were calciphiles
including Aesculus pavia, Elymus virginiana, Mat
and Yucca aloifolia.

The most inclusive study of the vascular flora at Indian middens includ-
ing shell mounds and rings is that by Dorroh (1971) along the South Caro-
lina coast. A shell mound is a solid mass of shellfish remains; and a shell ring
is a circular or arc-shaped ridge consisting of shellfish remains. Dorroh sampled
eleven mounds and eight shell rings during the summer of 1970 by transects.
At each site, a north south and east west compass line was followed, and
species were identified if they were encountered at the transect and approximately
two yards on either side of the transect. Dorroh’s (1971) transect approach
yielded 136 species in 59 families. The present study, a comprehensive in-
ventory of 9 shell rings and one shell mound, yielded 241 species.

There are several archeological surveys of shell rings, including one by a
future president, William McKinley, who described and measured three shell
rings at Sapelo Island, Georgia, for the Smithsonian Institution in 1872
(Calmes 1967). Waring and Larson (1968) studied the shell rings at Sapelo
Island and Porcher’s Bluff, South Carolina, just north of Charleston. Cameron
(1976) presented an ethnobotanical and floristic reconstruction of the Sapelo
Island Shell Ring. Dr. Warren Moorehead examined the Chesterfield site, a
horseshoe shaped structure along the Broad River near Port Royal Sound,
South Carolina, (Flannery 1943). The Sewee Ring, a horseshoe shaped midden
near the Santee River, South Carolina, was examined by Edwards (1965)
who postulated that the site may have served as a fish trap by native Ameri-
cans. Calmes (1967) examined the Ford’s Skull Creek and Sea Pines rings
at Hilton Head Island, South Carolina.

The Fig Island Shell Ring near the northeast end of Edisto Island, South
Carolina, was examined by Hemmings (1970). Trinkley (1980, 1985) has
done extensive work on the archeology of various shell rings and middens
in South Carolina including Buzzards Island, and Crow Island at the Francis
Porcher property north of Charleston.

Sageretia minutiflora,

METHODS
Nine shell rings and one shell mound were selected for the present study.
These include four rings in South Carolina: Sewee, Buzzards Island, Ford’s
Skull Creek, Sea Pines and the Auld Mound; three rings in Georgia: Oemler,

864 SipA 18(3)

Romerly Marsh and Sapelo Island; and two rings in Florida: Fort George
and Guana River State Park (Figure 1). Of the four shell rings extant in
Florida; two are in this study. In South Carolina there were over 100 shell
rings but many of these no longer exist due to shell removal for road build-
ing. Criteria for selecting sites include permission from private, state and
federal authorities, lack of human disturbance and accessibility. The study
was initiated in February 1993 and was completed in October 1994. Each
site was sampled a minimum of 6 times during the growing seasons of 1993
and 1994 for a total of about 40 field days. Herbarium vouchers of each
taxon were prepared and deposited at the University of South Carolina Herbarium:
some are also housed at the Brooklyn Botanic Garden (BEL), Missouri Botanical
Garden (MQ), and the New York Museum at Albany, N.Y. (NYS).

The species checklist (Appendix I) contains an inventory of the vascu

pascal

ar
plants that reproduce spontaneously and persist for more than one year without
cultivation, including native taxa, naturalized and adventive weeds, and escapes
from cultivation. The checklist is divided into the following categories: vascular
cryptograms, gymnosperms, dicots, and monocots. Nomenclature follows
Kartesz (1994); when differences occur, the name as presented in Radford
et al. (1968) is listed as a synonym and enclosed in brackets. The concept of
families follows Kartesz (1994).

Mineral analysis of the soils of the shell rings was provided by the Cornell
Nutrient Analysis Laboratory, Cornell University.

RESULTS AND DISCUSSION

The vascular flora of ten shell sites (9 rings, | shell mound) from South Carolina

to Florida consists of 80 families, 182 genera, and 241 species of which 216

(89.6%) are native (Table 1). There is a higher percentage (89.6%) of na-

tive flora at the shell middens than at Fort Moultrie, South Carolina, where

77% of the flora are native (Stalter & Lamont 1993). The native flora at the

Outer Banks of North Carolina, Ocracoke Island to the Virginia border, is
78% (Stalter & Lamont 1997).

The Poaceae, with 30 genera and 49 species and the Asteraceae, with 24
genera and 29 species are the largest families in terms of taxa. Together they
comprise 30% of all genera and 33% of all species. Other large families are
the Fabaceae and Cyperaceae. The largest genus is Dichanthelium with 8 species
followed by Overcus with 6 species. The rarest species encountered is Sageretia
minutiflova (Radford et al. 1971), though this species is common at shell
ring sites examined in the present study. A summary of the vascular flora is
given in Table

The Sewee Shell Ring, South Carolina, was visited by Dr. R. Mohlenbrock
in 1979. Notable species on Mohlenbrock’s list at the Sewee Shell Ring included
small-leaved buc an (Bumelia tenax) and basswood (Tilia caroliniana).

STALTER ET AL., Flora of Indian shell rings 865

Taste 1. A summary of the vascular flora identified at ten shell sites from South Carolina to Florida.
Ferns Conifers Dicots Monocots Total
Families 2 2 63 13 80
Genera 2 2 129 49 182
Spec 2 Z 164 13 241
oe i 2 2 142 62 208
Non-native oe 0 0 20 6 26

Shumard’s oak (Quercus shumardii), as well as climbing hydrangea (Ducumeria
barbara), grew only in the nearby forest. The forest was leveled by Hurri-
cane Hugo, September 21 and 22, 1989, and was severely burned by a dev-
astating wildfire in 1991. The vegetation on the shell ring escaped the fire,
but not the powerful winds of Hurricane Hugo.

Several factors may account for the assemblage of species on the shell sites
at the time this study was conducted. Hurricane Hugo, September 1989,
savaged three shell middens: Sewee, Auld and Buzzard’s Island, South Carolina.
Four study sites, Auld, Buzzard’s Island, Roemerly Marsh and the Oemler
Ring are islands saaroundled by a sea of Spartina alterniflora. Two shell rings,
Sewee and Sapelo Island border salt marshes; the vegetation of these two
rings and the four previously mentioned island rings are strongly influenced
by tidal inundation. Sea Pines, South Carolina, Ford Skull Creek, South Carolina,
Fort George and Guana River State Park, Florida are inland rings not influenced
by tidal flooding. The calcareous nature of the soil also affects species dis-
tribution. Human activity (disturbance) in the form of occasional mowing
at Sea Pines, archaeological excavation by Waring and Larson (1968) at Sapelo
Island and light cattle grazing at Sapelo Island, play a role in species distri-
bution at these shell rings. A final obvious factor is climatic differences.
The winters at St. Augustine, Florida, are milder than the winter climate
at the most northern Sewee Shell Ring site.

Sewee, Auld and Buzzards Island, South Carolina, were ravaged by Hur-
ricane Hugo, September 21 and 22, 9. Hurricane Hugo, a category 4
hurricane, was one of the most powerful storms of this century to strike the
South Carolina coast. The hurricane surge in the vicinity of Charleston, South
Carolina, was 4m; the surge reached 6m at McClellenville, 5Okm northeast
of Charleston. Hurricanes with winds as severe as those of Hugo strike por-
tions of the South Carolina coast approximately once every 200 years (Anonymous
1974). The vegetation on the above three shell rings was severely damaged
by wind. Additional damage to shell ring vegetation may have occurred in
one or more of the following ways: inundation by the storm surge; salt water
immersion and residual effect of salt water desiccation; smothering by deposition
of soil and rafts of vegetation (Stalter & Lamont 1993)

866 Sipa 18(3)

—
~

Stalter and Lamont (1993) recorded 218 species at Fort Moultrie, Sout
Carolina, a site just north of Charleston, one year after Hurricane Hugo,
including coastal plants, e.g. Quercus virginiana, Tex vomitoria, and Juniperus
silicicola that are common along the southeast coast. Stalter and Lamont (1993)
reported that many of the plants at Fort Moultrie and Fort Sumter survived
Hurricane Hugo. Liriodendron tulipifera and Podophyllum peltatum, two spe-
cies recorded by Gregory (1925) and Dorrah (1971) at the Auld Shell site
were probably killed by Hurricane Hugo. Yet many species identified by
Dorroh (1971) at the Auld site, e.g. Quercus virginiana, Smilax spp. survived
Hurricane Hugo.

The Sea Pines, South Carolina Shell Ring is maintained by occasional
mowing as an “archeological show piece” of the Sea Pines Development
community. Common shrubs, e.g. Myrica cerifera and Quercus virginiana and
saplings of Quercus laurifolia, abundant in the surrounding forest, have been
removed from this shell ring. Dichanthelinm spp., Oplismenus setarius and Arisaema
spp. provide conspicuous seasonal ground cover at this site in the absence of
dense shrub cover. The nearby Ford Skull Creek Shell Ring, a historically
“disturbed” inland shell ring, was once used as a source of oyster shells, as
building materials. At the time the present study was conducted, the Ford
Skull Creek Shell Ring was undisturbed. Past oyster shell harvesting may
have an effect on the present floristic composition at this shell ring.

The Sapelo Island, Georgia, Shell Ring, the most floristically rich shell
ring of this study, has been disturbed in the past. Waring and Larson (1968)
excavated shells from a portion of this shell ring bordering the salt marsh.
Today, feral cattle occasionally graze portions of the shell ring, though grazing
appeared to be negligible at the time of the present study. The presence of
exclosures at this shell ring, might enable future investigators to determine
the impact of cattle grazing on the flora of this shell ring.

Two Florida shell rings, Fort George, 23km east of Jacksonville and Guana
River State Park, just north of St. Augustine, are undisturbed inland shell
rings. Vernonia gigantea, Psychotria nervosa, Peperomia humilis and Bidens alba
var. radiata were found only at the Florida shell rings; they were not ob-
served at the other shell rings. Coral/lorhiza wisteriana was observed in flower
in mid February, 1994, at the Guana River site, while at Fort George, 5Skm
north, C. wzsteriana did not flower until late March, 1994. Yet, the climatic
differences between the two sites are minimal (Anonymous 1974). The dif-
ference in flowering of C. wisteriana at the two sites may reflect significance
differences in minimum temperature at these sites during the winter of 1994.

Eleuterius and Otvos (1979) report that Juniperus silicicola, Aesculus pavia,
Erythrina herbacea, and Morus rubra are reliable indicators of the shell de-
posits on Indian middens. All of the aforementioned species were found at

STALTER ET AL., Flora of Indian shell rings 867

one or more of the shell rings in the present study. Sageretia minutiflora and
Hexalectris spicata may also be included as “indicator species” as these spe-
cies favor the calcareous soils of Indian middens (Wunderlin 1998) but are
found at non-ring sites as well.

The soils at the shell rings are neutral to slightly basic with a pH range
of 6.82 at Sea Pines to 7.69 at Guana River State Park. Available Ca was
high at all sites with a range of 1497 mg/Kg at Buzzard Island to 29,706
mg/Kg at Sea Pines. Available Mg ranged from 139.9 mg/Kg at Sapelo to
921.1 at Buzzards Island, while available Mn ranged from 5.4 at Fort George
to 62.5 mg/Kg at Sea Pines. Available P ranged from 19.2 at Fort George
to 221.0 mg/Kg at Sapelo, while K ranged from 42 at Guana River to 207
mg/Kg at Buzzards Island. Available nitrates ranged from undetectable at
Buzzards Island to 190.61 at Sea Pines. The range of all these nutrients is
highly variable among the sites.

Several shell rings occur as islands in salt marshes. The distribution of
salt marsh species on the sides of these rings is related to tidal flooding and
soil salinity. Spartina alterniflora occupies the lowest daily flooded base of
the shell ring. Less flood tolerant salt marsh species are Sa/icornia spp., Batis
maritima, Distichlis spicata, Borrichia frutescens, and Spartina patens. These form
distinct zones above the more flood tolerant Spartina alterniflora. Iva frutescens
and Baccharis halimifolia border Borrichia and Spartina patens, on the upland
side. By contrast, the vegetation on shell rings not bordering salt marshes
includes Arisaema dracontinm and A. triphyllum at Sea Pines and Ford’s Skull
Creek, South Carolina, while Cora/lorhiza wisteriana and Hexalectris spicata
were observed at the two Florida shell rings. Psychotria nervosa, Peperomia
humilis and Bidens alba var. radiata, three subtropical species, were exclu-
sively at the Florida shell ring sites, and are not part of the Carolina flora.

The most common woody species found on the salt marsh island shell
rings are Juniperus silicicola, Quercus virginiana, Sabal palmetto, [lex vomitoria,
Myrica cerifera and Smilax spp. The most notable plant records, exclusive of the
rare Sageretia minutiflora, ace Liriodendron tulipifera, Podophyllum peltatum, and
Trillium sp., known only from historical records at the Auld South Carolina Shel
site (Gregorie 1925). Dorroh (1971) reported Lirtodendron and Podophyllum
at Auld in her study. These aforementioned taxa are usually not found at coastal
barrier islands, although Stalter and Lamont (1987) reported a small popu-
lation of Podophyllum peltatum and a single Liriodendron tulipifera at Assateague
Island, Virginia. Liriodendron, Podophyllum and Trillium were absent at the Isle
of Palms, South Carolina (Stalter 1976), Turtle Island, South Carolina (Stalter
1973), coastal Brookgreen Gardens, South Carolina (Stalter 1972), the Outer
Banks of North Carolina (Stalter and Lamont 1997), and Fisherman Island,
Virginia (Stalter et al. 1997). The Liriodendron at the Auld Shell mound

oes

868 Sipa 18(3)

may have been toppled by Hugo’s hurricane winds while Podophyllum may
have been killed by saltwater inundation during the storm surge associated
with Hurricane Hugo, September 1989. Radford et al. (1971) report that
Liriodendron, Podophyllum and Trillium are present at Charleston County, South
Carolina, though these taxa may have been collected at inland sites.

Many plants may be “rare”, rare being defined as scarce, less than 5 populations,
at a study site (Stalter & Lamont 1997). Scalter and Lamont (1997) recorded
336 “rare” plants, 45.3% of the flora, at the Outer Banks of North Carolina.

Plants are rare for a multiplicity of reasons. Plants may be rare because
they exist at the edge of their normal range. Plants may be rare if they have
narrow habitat requirements that are met in only a few areas, such as the
calciphile, Sageretia minutiflora. Plant succession, climatic events such as drought,

—

severe cold or hurricanes can reduce the number of species, especially plants
in rescricted habitats such as shell rings. Human activities such as develop-
ment, introduction of non-native plants, pollution and overcollection of attractive
plants such as Cora/lorhiza wisteriana may also contribute to the rarity of
species (Stalter & Lamont 1998).

Sea level along the Atlantic coast was 120 meters lower than today 12,000
years ago. Dolan et al. (1980) report that sea level continued to rise for 8,000
years, ‘reaching within afew meters of the present level 4-S000 years ago.”
Sea level has risen several meters in the past 2,000 years, and over the past
100 years, sea level has risen over 30 cm (Dolan et al. 1980). It is possible
that some of the present salt marsh island shell rings may have been ini-
tially located on upland sites when they were built by Native Americans,
several thousand years ago, when the sea level was several meters lower than
it is today (Dolan et al. 1980). All the present shell ring sites selected in
this study may have been built close to, or on the coast, where oysters were
abundant. However, the precise location of the shell rings examined in this
study relative to the coastline, salt marsh creeks and/or salt marshes at the
time of their construction cannot be accurately determined.

APPENDIX I
ANNOTATED CHECKLIST OF SPECIES

The vascular plant taxa found at ten shell sites have been arranged ac-
cording to the following categories: vascular cryptogams, gymnosperms,
dicots, and monocots. Within each category, families and lower taxa are arranged
alphabetically. Nomenclature primarily follows Kartesz (1994).

Each entry includes the following information sequence: scientific name;
pertinent synonym, enclosed in brackets; and frequency relative to the shell
rings, using the categories: rare (scarce, less than 5 populations), infrequent
(uncommon, occasional, 5 to 20 populations), frequent (common, more than
20 populations).

STALTER ET AL., Flora of Indian shell rings

POLYPODIOPHYTA

OLYPOD
Teas ces (L.) Andrews and
Windham spp. michauxiana (Weatherby)
Andrews & Windham
pol: wpodivide de)

eat one infrequent

Polypodium

Watt. var. 216) 7h) tUXIANH I

ASPLENIACE
Aspleninm a neuron (L.) BSP.; frequent
PINOPHYTA
CUPRESSACEAE
Juniperus virginiana L. var. silicicola (Small)
E. Murray [Juniperus silicicola (Small)
Bailey}; infrequent
PINACEAE
Pinus taeda L.;

MAGNOLIOPHYTA
MAGNOLIOPSIDA (DICOTS)

infrequent

SANTHACEAE
Ruellia caroliniensis J.F. Gmel.) Steudel;
seas

ACER :
Acer ee L.; infrequent
AMARANTHACEAE

Tresine rhizomatosa Standley; rare

ACARDIACEAE
Toxicodendron radicans - ) Kuntze [Rhus

radicans L}, frequ
Rhus alvin Ly ae copallina L.J;
infreque

ONACEAE
Asimina triloba (1...) Dunal;: rare
APIACEAE
Sanicula canadensis L.:
AQUIFOLIACEAE
Ilex vomitoria Aiton, infrequent

infrequent

ARALIACEAE
Aralia spinosa L.;
ASCLEPIADACEAE

Cynanchum angustifolinm Pers.; {Cynanchum
palustre (Pursh) Heller]; infrequent
Matelea gonocarpa (Walter) Shinners [M.
suberosa (L.) Shinners}; rare

rare

869

ASTERACEAE
Ambrosia artemtsitfolia L.; rare
Arnoglossum ovatum (Walt.) H.E. Robins.
{[Cacalia lanceolata Nutt. - rare
Aster tenuifolinus L.; infreq
Baccharis muniiale Michx.: ; rare
Baccharis halimifolia L.; frequent
ge Ms (L.) DC. var. — ultz-
p.) Ballard ex T.E. Melchert es
oe L. var. radiata Schultz — Bip.};
frequent at Florida Shell Rings
Bidens bipinnata L.; infrequent
Borrichia frutescens (L.) DC.; frequent
Carduus smallii (Britton) Ahles. [Czrsiam
horridulum Michx.}; rare
Leacanthemum vulgare Lam. {Chrysanthemum
leucanthemum J; infrequent
Conyza canadensis (L.) Cronq. var. pusilla
Cronq.; infrequent
al ol tomentosus L.; frequent
Erechtites =
Erigeron quert vifoli reque
Eupatorium ieee — ) Gail
reque
Eupatorium byssopifolium L.; Lots
Eupatorium serotinum Niche
Euthamia alae (L.)N vt, (Solidago
graminifolia (L.)S nea infrequent
Gamochaeta be ee Ge '
[Gnaphalium purpureum L. var. purpurenm);
infrequent
Hieracium gronovii L.;
Iva frutescens L.; infrequent
Lac ae va (L.) Gaertn.; infrequent
Lactuca graminifolia ee ; infrequent
Mikania scandens (L.) Willd.; frequent
Smatlanthus nvedalia (L.) ee ex Small
[Polymnia uvedalia (L.) L.}; rare
Solidago sempervirens L. var. mexicana (L.)

infrequent

Fern.; frequent
*Sonchus asper (L.) Hill; frequent
Verbesina occidentalis (L.) Walter; rare
Vernonia gigantea (Walt.) Trel.; ra
BATACEAE

Batis maritima 1.; rare

BERBER
Paap beltatun L. Reported by
e (1925) and Dorroh (1971) at
rein not recently observed

870

a

BIGNONIACEAE
Rj 2 capveotata lL. £ TA nisostichus cat

(L.) Bureau]; infrequent
Campsis radicans (L.) Seemann; infrequent
BORAGINACEAE
Heliotropinm curassavicum L.; rare

BRASSICACEAE

*Cardamine hirsuta L.;
Descurainia pinnata (Walter) Britton ssp.

brachycarpa (Richardson) Delting [D.
1) O.E. Schultz};

infrequent

7 | ]
brachycarpa Richard

rare
Lepidium densiflorum Schader; rare
— virginicum L.; frequent
BUDDLEJACEAE
Pol) pean procumbens L.; infrequent

CACTACEAE
Opuntia humifusa Rat.) Rat. [O. compressa
J.E. Macbride}; infrequent

*OQpuntia monacantha (Willd.) Haw. [0.
— Mill. dsr
PANULACEAE

ee (L.) Nieuwl. [Specalaria
perfoliata (L.) A. DC.1J, frequent

CAPRIFOLIACEAE
*Lonicera japonica Thunb.;
Lonicera sempervirens L.; infrequent

rare

CAM

infrequent

Viburnum nudum L.;
CARYOPHYLLACEAE

Arenaria lanuginosa (Michx.) Rohrb.;

aca ia serpyllifolia L.; infrequent
Baumg. subsp. v/gare
eg rne Greuter & Burdet [C.
holosteotdes Fries var. vulgare (Hartman)

saad L.}; infrequent
ina L.; rar

> fare

WUD?

tray toe
J

Hylander; C.
Silene antirrhty
* Sher iaite marina (L. ’ Griseb

eK. Presi}; infrequent

LS. salina

CELASTRACI
Enonymus americanus L.; rare
CHENOPODIACEAE
Atriplex patula 1..; infrequent
*Chenopodium album L.; infrequent
*Chenopodinm ambrosioides L.; frequent

*Salicornia maritima Wolff & Jefferies
auct.non L.];

Salicornia

a

enropaed

infrequent

Sipa 18(3)

frequent
infrequent

Salicornia virginica L.;
Suaeda linearis (Elliott) Mogq.;
CONVOLVULACEAE
Dichondra carolinensis Michx.; infrequent
Ipomoea pandurata (L.) G.F.W. Mey.;
infrequent
[Ipomoea sagittata Poir.; frequent
CORNACEAE
Cornus asperifolia Michx.; rare
Cornus florida L.; rare
Cornus stricta Lam. {C. foemina P. Mil
infrequent

|;

CUSCUTACEAE (F pets Convolvulaceae)
Cuscuta gronovit Willd.;
ERICACEAE
Vaccinium corymbosum L. {V. atrococcum (A.
Gray)
EUPHORBIACEAE
Acalypha gracilens A. Gray; rare
eee maculata (L.) Small {Euphor-
bia maculata L.; E. supina Raf.}; infrequent
C pee se Nes (Michx.) Engelm. &
ray; frequent
E bee hia cyathophora Murray {E. heterophylla
L. var. cyathophora (Murray) Griseb.}; rare

orter]}; rare

FABACEAE
Cassia nictitans L.; a
Cercis canadensis L.; rare

Clitoria mariana ie

C rotolari ld iiniaae Wal ex. J.E Gmel.

_ angilata auct. Mill. = rare

De. smodinum nndiflran aL. ) a
*D—)

=
~

eer

ae beibenes ie Fecuien
L.) BSP.; infrequent
Lespedeza sp.; infrequent
Lespedeza stuevei Nutt.; infrequent
ees officinalis (L.) Lam.; se
Visterta sinensis (Sims) DC.,;

Galactia regularts (

FAGACEAI
Quercus laurifolia Michx.; rare
Quercus ie ae Willd.; rare

Quercus nigra L.

Quercus phellos ie re
Quercus stellata Wangeoh.;
Quercus virginiana Miller; frequent

rare

GERANIACEAE

Geranium carolinianum L.; infrequent

STALTER ET AL., Flora of Indian shell rings
HAMAMELIDACEAE

Hamamelis virginiana L.; rare

Liquidambar styracifiua L.; rare
HIPPOCASTANACEAE

Aesculus pavia L.; frequent

JUGI ANDACEAE

Carya glabra (P. Mill.) Sweet; infrequent
Juglans nigra L.; rare
LAMIACEAE
Salvia lyrata L.; frequent
Teucrium canadense L.; frequent
Trichostema dichotomum 1..;
LAURACEAE
se Sed radi . ) Sprengel;
idum (Nutt.) Nees
LOGANIACEAE
Gelsemium
infrequent

rare

infrequent
. infrequent

sempervirens (L.) St.-Hil.;
Polypremum procumbens L.; infrequent
Spieelta marilandica L.; ra
MAGNOLIACEAE
Liriodendron tulipifera L. Reported by
Gregorie (1925) and Dorroh (1971) at
Au Id, SC. Probably destroyed by Hur-
ne Hugo in 1989

ia grandiflora L.; sabequent

Magnolia virginiana L.; infrequent
MELIACEAE
* Melia azedarach L.; rare

MENISPERMACEAE

Cocculus carolinus (L.) DC.; rare

Menispermum canadense L.; rare
MORACEAE

* Morus alba L.; rare

Morus rubra L.; infrequent
MYRICACEAE

Myrica cerifera L.; frequent
NYSSACEAE

Nyssa sylvatica Marsh.; infrequen

Nyssa biflora Walt. {N. sylvatica Marsh. var.
biflora (Walt.) Sarg.}; infrequent

OLEACEAE

Fraxinus cavoliniana P. Mill.; rare
*Ligustrum amurense Carr., infrequent
Osmanthus americanus (L.) A. Gray;
infrequen

NAGRACEAE

Ocnothera fruticosa L.; frequent

Ocnothera laciniata Hill, infrequent
OXALIDACEAE

Oxalis dillenii Jacq.; infrequent

Oxalis stricta L.; infrequent
PASSIFLORACEAE
Passiflora lutea L.; infrequent

PHRYMACEAE

Phryma leptostachya L.; rare
PHYTOLACCACEAE

Phytolacca americana L.; frequent
PIPERACEAE

Peperomia humilis A. Dietr.; rare

PLUMBAGINACEAE
Limonium carolintanum (Walt.) Britt. {L.
nashit Small); infrequent
POLYGON ACEAE
Rumex hastalalus

Baldw.; frequent
RANUNCULACEAE
Clematis crispa L.; infrequent
*Clemantis terniflora DC. {Clematis
lioscoreifolia Lev\. and Vaniot}; infrequent
€

Clematis virginiana L.; ra
RHAMNACEAE
Berchemia scandens (Hill) K. Koch; infrequent
Frangula caroliniana (Walt.) Gray [Rhamnus
niana Walt.}; rare
Sageretia minutiflora (Michx.) C. Mohr
[Rhamnus minutiflora Michx.}; infrequent
ROSACEAE
Crateagus uniflora Muenchh.
Prunus caroliniana (P. Mill.) Ait. ioe
carolinina (P. Mill.) M. Roemer];
infrequent
Prunus serotina Ehrht.;
Rubus trivialis Michx.;
RUBIACEAE
Galium pilosum Aiton; infrequent

carolin

infrequent
frequent

Psychotria nervosa Sw. {Psychotria undata
Jacq.}; rare
RUTACEAE
Zanthoxylum clava-herculis L.; infrequent
SAPOTACEAE
Sider aaa tenax L. {Bumelia tenax (L.)
Willd.}; infrequent

872

SCROPHULARIACEAE
Gratiola pilosa Michx.; rare
Nuttallanthus canadensts (L.) D.A. Sutton.
na? is (L.) Chaz.}, frequent
*Verbascum thapsus L.; frequent
*Verontca arvensis L.; frequent
SOLANACEAE
Physalis walteri Nutt. {P. viscosa L. subsp.
maritima (M. A. Curtis) Waterfall];
infrequent
TILIACEAI
Tilia americana L. var. caroliniana (P. Mill.)
Castigl. [T. caroliniana P. Mill); rare
ULMACEAE
Celtis laevigata Willd.; infrequent
Celtis tenuifolia Nutt. [C. oxidentalis L. var.
georgiana (Small) Ahles}; rare
VERBENACEAE
Callicarpa americana L.;
Verbena scabra Vahl; ra
VIOLACEAE
Viola sororia Willd.;
VITACEAE

A mpelof bere arbored

‘ta canadens

in —— uent

rare

L.) Koehne; infrequent
Pusthinciins spre (L.)) Planch,
frec
Vitis ok Michx.; infrequent
Vitis rotundifolia Michx.; infrequent
MAGNOLIOPHYTA -
LILIOPSIDA
AGAVACEAE
*Vncca — Le
ARACEA
yea od ‘acontinm (L..) Schott; infrequent
Arisaema triphyllum (L.) Schott;
ARECACEAE

Sabal palmetto Lodd. ex. Schultes; infrequent

infrequent

rare

BROMELIACEAI

Tillandsia usneoides (L.) L.; infrequent

COMMELINACEAE

Tradescantia obiensis Raf.; infrequent
CYPERACEA
Cladium jamaicense Crantz [C. mariscus (L.)
Pohl}; infrequent
Cyperus globutosus Aublet; infrequent

Sipa 18(3)
Cyperus retrorsus Chapm.;
Eleocharis albida Vorr.; rare
Fimbristylts castanea (Michx.) Vahl, infre-
quent
ee eh colorata (L.) H. Pfeiffer
hromena tos L.) Hitchce.}; rare

infrequent

oe robustus Pursh; rare
Scleria mes rata Michx. {S. vitida Muhl.};
infrequent

DIOSCOREACEAE
Dioscorea villosa L.;
IRIDACEAE
Sisyrinchium rosulatum E. Bickn.; rare
JUNCACEAE
Juncus bufonins L.;
Juncus roemertanus Scheel
LILIACEAE
*A lium vineale L

rare

infrequent

e; infrequent

L.; infrequent
Trillium sp. Reported by Gregorie (1925)
t Auld

ORCHIDACEAE

site; not recently observed.

Corallorhiza wisteriana Conrad; infrequent
Hexalectris spicata (Walt.) Barnh.; infrequent
POACEAE
And) aan elomeratus (Walter) B.S.P. [A.
vir, ginicus L. var. abbreviatus (Hackel

Fern.

—S

m}; infrequent

Andropogon virginicus L.; frequent

Arundinaria gigantea (Walter) Muhl. subsp.
tecta (Walter) McClure; infrequent

Arthraxon hispidus Thunb. var. lala
(Hackel) Pipnea ee

*Bromus tectorum L., infrequent

Cenchrus carolinianus Walt. {C. meertus M.A.
Curtis}; rare

Chasmanthinm laxum (1..) Yates (Uniola laxa
(L.) B.S.P.}; infrequent

*Dactyloctenium aegyptinm (L.) Willd.; in-
frequ

ig rea acuminatum (Sw.) Gould &

k [Panicum acuminatum Sw. (sensu

oe incl. P awburne Ashe; P. meridtonatle

Ashe; P. lewcothrix Nash}, infrequent
Py oar fos besenie ; (1 | Ba ae

Dj;-/ 1]

& Clark {Panicum consanguineum Kunth};
infrequent

Dichanthelinm latifolinm (.) Gould & C.A.
Clark [Panicum latifolinm L.}; infrequent

STALTER ET AL., Flora of Indian shell rings 873

Dichanthelium laxiflorum (Lam.) Gould Panicum OEE Michx.; infrequent
{Panicum laxiflorum Lam.; P. xalapense Panicum virgatum L.; infrequent
.}; rare Paspalum distichum L.; amas
Dichanthelt bhyllum (Nash) Gould Paspalum setaceum Mic tne ee
[Panicum malacaph) Lum pa rare ichx.) Vasey [P. viliatifolium Mic
bantheltt sosanthes var. scriberanum P. lon eae LeConte}]; rare
(Nash) Coad oer em Pid: caroliniana Walt.; rare
Nash}; rare Piptochaetium avenaceum (L.) Parodi {Stipa
Dichanthelinm sabulorum (Lam.) Gould & avenacea L.]; infrequent
Clark var. patulum (Scribn. & Merr.) *Poa annua L.; infrequent
Gould & Clark. [Panicum lancearinm *Polypogon ee (L.) Desf.; frequent
Trin.]; infrequent Setaria genicilata (Lam.) Beauv.; infrequent
Dichanthelinm scabrinsculum (Ell.) Gould Sp oe oe (Rydb.) Rydb.; rare
Clark. [P m scabriusculum Ell. Sphenopholis obtusata (Michx. es rare
actleatum A. Hite che. & Chase}; rare _ alter sls Loisel.; frequ
Distichlis spicata (L.) Gecae infrequent Spar tens (Ait.) Muhl. nnn
Echinochloa walteri (Pursh) Heller; rare ance: indicus a )R. Br. [s. poiretti(R.
Elymus virginicus L. var. halophilus (Bickn.) & $.) Hitchc.}; frequent
Wieg.; infrequent Sporobolus virginicus (L.) Kunth; infrequent
Eragrostis hirsuta (Michx.) Nees; infrequent Stenotaphrum secundatum (Walt.) Kuntze;
Eragrostis spectabilis (Pursh) Steud. infrequent
infrequent Tridens flavus (L.) A. Hitche. [Trivdia flava
Eustachys petraea pee Desv. [Chloris (L.) Smyth}; infrequent
iis Sw.]}; infrequent Triplasis purpurea (Walt.) Chapman;
Melica mutica Walt. a ane requent
Mable ee ae Chath.) Tein: Vulpia octoflora (alt.) Rydb. [Festuca octoflora
Wale.]; infrequent
ico Cada EF. Gmel.; rare SMILACACEAE
oe eae ) Roemer & Schulte: Smilax bona-nox L.; infrequent
lus (L.) Beauv. subsp. setarius Smilax glauca Walt.; rare
(aa Mer ex Ekman]; frequent milax ia uvifolia L.; infrequent
Panicum amarum E\L. {P. amarnlum A. Hitchc. Smilax rotundifolia L.; infrequent

Chase}; rare
pies boscit Poir. {P. bosciz var. molle oe
itchc. and Chase}; infreque

ACKNOWLEDGMENTS

For access to the following shell ring sites we thank: Mr. Francis Porcher
(Buzzard’s Island), Sea Pines Corporation (Sea Pines), Mr. Ford (Ford’s Skull
Creek), Mr. Ralph Hinz, The Landings (Romerly Marsh), University of Georgia
Marine Institute (Sapelo Island), Mark Epstein, The Florida Game and Fresh
Water Fish Commission (Fort George and Guana River). Thanks are also
extended to the University of Georgia for providing me with transporta-
tion to the Sapelo Island Shell Ring, and to Chester dePratter and Michael
Trinkley for providing directions to several shell rings. For mineral analy-
sis, we thank the Cornell Nutrient Analysis Laboratory. For field assistance,
we thank John Baden and Paul Teller. For assistance in the identification of

several taxa we thank Ihsan Al-Shehbaz, Steven Clemants, Robert Meyer,
Richard Mitchell, James Montgomery, Richard Rabeler, Charles Sheviak,
and Gordon Tucker. For reviewing the paper, we thank Steve Dial. Finally, we
acknowledge the assistance of Michelle Bailey, undergraduate research stu-
dent at St. John’s University, and the financial support of St. John’s University.

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Brown, C.A. 1936. The ve seni one Indian mounds, middens, and marshes in Plaquemines
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Catmes, A. 1967. Test excavations at Two Late Archaic Sites on Hilton Head Island. Ms.
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CAMERON, M.A. 19 76. Sapelo Island: An ethnobotanical and floristic reconstruction, Un-
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9

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DePrarrer, C.B. 1974. An en ee survey ee saw Island, Chatham County, Georgia:
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Epwarps, W.E. 1965. Preliminary report on the Sewee Indi
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lumbia.

ELeurerius, L.N. and L.G. Orvos 1979. Floristic and geologic aspects of Indian mid
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FLANNERY, R. 1943. Some notes on a few sites in Beaufort County, South Carolina, Bull.
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Forp, J.A 1966. E ‘arly formative cultures in Georgia and Florida. Amer. Antiquity 31:781—
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ens

GREGORIE, A.K. 1925. Notes on Sewee Indians and Indian remains of Christ Church Par-
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Hemmincs, E.T. 1970. Emergence of formative life on the Atlantic Coast of the Southeast
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876 Sipa L8(34)

BOOKS RECEIVED

Fautin, DapHNe Gait, DouGtas J. FuruymMa, and FRANcEs C. JAMES, eds.
1998. Annual Review of Ecology and Systematics. Volume 29.
(ISBN 0-8243-1427-1, hbk). Annual Reviews Inc., 4139 E] Camino
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. Individuals: $60.00
USA, $65.00 Elsewhere; Institutional: $120.00 USA, $130.00 Else-
where. 626 pp.

Volume 29 of Annual Review of Ecology and Sy stematics is another excellent mix of articles
on ecology and systematics. A total of 20 articles are presented followed by a Subject In-
dex, Cumulative Index of Contributing es anda ciate: Index of Chapter Titles
Volumes 25—29
Contents include:

Molecular Trans-Species Polymorphis

Principles of Phylogeography as ener by Freshwater and Terrestrial Turtles in the

Southeastern United States

The Functional Significance of the Hyporheic Zone in Streams and Rivers

Endangered Mutualisms: The Conservation of Plant-Pollinator Interactic

The Role of Introduced Species in the Degradation of Island Ecosystems: Case His-

tory of Gua

Evolution of Helping Behavior in Cooperatively Breeding Birds

The Ecological Evolution of Reefs

Roads and Their ee Ecological Effects

ation ios, and Genetic Conflict
Early Evolution of Land Biante: Phylogeny, Physiology, and Ecology of the Primary Terrestrial
Radiation
Possible Largest-Scale Trends in Se rane Evolution: Eight “Live Hypotheses”
Fungal Endophytes: A Continuum of Interactions with Host Plants
Floral Symmetry and Its Role in Plant- ia Systems: Termino

ss

ogy, Distribution,

and Hypotheses
Vertebrate Brae in Marine and Terrestrial Environments: A Nutritiona

Ecology
Perspective
Carbon and Carbonate oo in Coastal Aquatic Ecosystems
e Scientific Basis of Forestry
ae s, Mechanisms, and Rates of Polyploid Formation in F
aa Growth Efficiencies in Natural Aquatic Systems
he Chemical Cycle and Bioaccumulation of Mercury

owering Plants

Th
Phylogeny of Vascular Plants

Stipa 18(3): 876. 1999

NOTEWORTHY VASCULAR PLANTS
FROM ARKANSAS

ERIC SUNDELL
School of Mathematical and Natural Sciences, University of Arkansas
Monticello, AR 71656, U.S.A
R. DALE THOMAS
Department of Biology, Northeast Louisiana University
Monroe, LA 71209, U.S.A.
CARL AMASON
P.O. Box 164,
Calion, AR 71724, U.S.A.
ROBERT L. STUCKEY
Rt. 1, Box 1440
Texarkana, AR 75502, U.S.A.
JOHN LOGAN
Arkansas Natural Heritage Commission
323 Center Street, Little Rock, AR 72201, U.S.A.
ABSTRACT

The authors provide a list of 41 additions, reinstatements, and significant range exten-
sions for the flora of Arkansas. Trautvetteria car oliniensss, Datlea gattingeri, Rhynchosia minima,
and Setaria pumila are reported as new and persistent elements, while Magnolia macrophylla,
oe diandra, Valerianella locusta, ee oes exile are reinstated to the state flora.

nun of alien weeds are noticed and documented for the first time, however, their

Se is noc know
RESUMEN

Los autores proporcionan una lista donde se enumeran 41 adiciones, confirmaciones, y
extensiones significativas para la flora de Arkansas. Trautvetteria caroliniensis, Dale see,

Rhynchosta minima, y Setaria pumila se citan como Componentes nuevos y persistente
drea, mientras que Magnolia macrophylla, Didiplis diandra, Valer sn locusta, y | Sisyrinchium
exile se confirman para la flora del estado. Un ntimero de malas hierbas de origen aldctono

cida su persistencia

son mencionadas y documentadas por primera vez, sin embargo, n
The authors provide a list of 41 taxa representing additions and rein-

statements to the Arkansas flora as well as noteworthy range extensions within
the state. Herbarium abbreviations are taken from Holmgren et al. (1990).

SIDA 18(3): 877-887. 1999

878 Sipa 18(3)

APIACEAE

Hydrocotyle sibthorpioides Lam. This invasive Asian pennywort occurs
sporadically in the Southeast in wet, disturbed habitats. We add three county
records to the two recorded by Smith (1988). Ashley Co.: Swndel/ 11,104
(UAM). Calhoun Co.: Thomas & Amason 157,168 (NLU, UAM). Pulaski
Co.: E. & M. Sundell 12,048 (UAM).

ASTERACEAE

Conyza bonariensis (L.) Crong. This common and widespread horseweed
has escaped notice, or at least collection, in southern Arkansas for years—
the more remarkable because it forms extensive populations along our major
highways that are easily recognized even at speeds of over 60 miles per hour.
Conyza bonariensis was not discovered in the state until 1989, when Thomas
and Amason collected plants ina railroad yard in E] Dorado 1n Union County
(Thomas et al. 1991). Voucher specimens taken from roadsides and river
banks are recorded here for eight additional counties. Arkansas Co.: Tho-
mas, Sundell, & Amason 145,284 (NLU). Calhoun Co.: Thomas & Amason
157,214 (NLU, UAM). Cleveland Co.: Thomas & Amason 157,108 (NLU,
UAM). Desha Co.: Thomas, Sundell, & Amason 145,498 (NLU). Drew Co.:
Sundell 12,110 (0AM). Hempstead Co.: Thomas & Amason 155,483 (NLU).
Lafayette Co.: Thomas, Sundell, & Amason 150,931, 156,555 (NLU). Miller
Co.: Thomas, Sundell, & Amason 151,143 (NLU). The species should be considered
an established element of the Arkansas flora.

Crepis setosa Haller f. is known from Missouri (Steyermark 1963) and the
northeastern United States (Gleason and Cronquist 1991) asa waif. Although
the species is unlikely to persist in Arkansas, its presence is documented by
a collection from Russellville in Pope County, where it grew as a weed in
pavement cracks, E., M., GJ. Sundell 12,105 (UAM).

BRASSICACEAE

Cardamine concatenata (Michx.) O. Schwartz. Like bloodroot and white
trout lily, toothwort is rare enough on Arkansas’ West Gulf Coastal Plain
to have gone undetected except for a single report for Arkansas County (Smith
1988). We record it here from mesic woodlands in two additional locali-
ties. Drew Co.: Swndell & Pagan 10,149 (UAM). Union Co.: Thomas & Amason
109,031 (NLU

Cardamine debilis D. Don is an unobtrusive and probably undercollected
alien weed, sporadically introduced in greenhouses and warmer areas of North
America (Rollins 1993). Plants were collected from a flower bed in Pulaski
County, October 1997, marking the species’ first occurrence in Arkansas.
Sundell 12,258 (UAM, UARK). Although Kartesz (1994) places C. debilis

SUNDELL ET AL., Vascular plants from Arkansas 879

in synonymy with C. flexwosa With., the two appear to be distinct and are
recognized as such by Rollins.

Cardamine flexuosa With. The alien Cardamine flexuosa grew in abundance
at Daylite Nursery in Drew County, where it had escaped from containers
to surrounding work areas. Rollins (1993) records this infrequently collected
cress from several eastern states, Mexico, and Costa Rica. Smith’s (1988)
report of the species from Baxter County, Arkansas, was based on misidentification
of material of Sibara virginica (Smith, pers. comm.). Thus we report it here

as new to Arkansas. Swndel/ 12,259 (NLU, UAM, UARK

Lepidium oblongum Small. Although Rollins (1993) describes the range
of this prostrate peppergrass as extending east to Arkansas, Smith (1988)
has seen only a single specimen from Lonoke County. We here add Wash-
ington County to that short list, where it was growing in pavement cracks
on the University of Arkansas campus in Fayetteville. Swndel/ 12,204 (UAM,
UARK). Collections at NLU document the species’ presence in Mississippi
and Louisiana as well.

Raphanus sativus L. Radish escapes from cultivation rather frequently in
the cooler parts of North America where it is common and abundant in
ruderal habitats (Rollins 1993). Although it apparently does not persist as
a weed in Arkansas, its occurrence should be noted. In Drew County, a few
plants were scattered along a newly seeded road construction site. Swnde//

12,260 (UAM

Rorippa sylvestris (L.) Besser. A collection from inside the Mississippi River
levee in Chicot County reconfirms the presence of this weedy crucifer in
Arkansas, previously reported for Crittenden County by Wilcox in 1973.
Sundell, Thomas, & Amason 10,952 (UAM).

COMMELINACEAE

Murdannia keisak (Hassk.) Hand.-Maz. This species has been known in
Arkansas only from Tucker’s 1969 report for Conway County. In 1996, Thomas
collected specimens on a shaded roadbank in nearby Pulaski County. T/o-
mas, Hunter, et al. 148,617 (NLU).

Murdannia nudiflora (L.) Brenan is an Asian introduction that has become
rather common in disturbed sites on the Atlantic and Gulf Coastal Plain
(Darwin et al. 1981, Wunderlin 1998). Thomas’s collection from Hempstead
County in southern Arkansas, where plants dominated a shaded lawn, rep-
resents its first record in the state. Thomas, Amason, et al. 155,570 (NLU).
Tradescantia crassula Link & Otto. Amason has observed this spiderwort

thriving out of doors at his home in Union County for several decades. Thomas,

Sundell, Amason, et al. 156,721 (NLU).

880 SIDA 18(3)

CYPERACEAE

Cyperus cuspidatus Kunth. This little flat sedge occurs sporadically on
the eastern Gulf Coastal Plain (Godfrey & Wooten 1979) and is recorded
here from two Arkansas counties. Chicot Co.: Thomas 142,804 (NLU). Drew
Co.: E. & J. Sundell 10,435 (UAM).

EUPHORBIACEAE

Phyllanthus pudens L.C. Wheeler and Phyllanthus tenellus Roxb. Both
Phyllanthus pudens (E. & M. Sundell 12,037 {UAM)) and P. tenellus (E. & M.
Sundell 12,038 [UAM}) were found thriving at Daylite Nursery in Drew
County where they were escaping to gardens and waste areas from plant
containers brought in from Louisiana and Texas. Although both of these
species reproduce out of doors in Ouachita Parish in north Louisiana, there
is no evidence that they will persist in the Arkansas flora. Phy/lanthus is
largely a tropical and subtropical group with only a single species, P. caroliniensis,
indigenous to the temperate regions of the southeastern United States.
Nevertheless, the recent, successful invasion of southern Arkansas by P. wrinaria,
first reported by Smith for Union and Arkansas counties (Smith 1978-1980)
and again by Sundell (1986) for additional localities, suggests that if trans-
portation is available, other species of Phy/lanthus might move north.

FABACEAE

Dalea gattingeri (Heller) Barneby. Known from cedar glades in Tennes-
see, Georgia, and Alabama (U[sely 1990) and, more recently, from Howell
County, Missouri, within 10 km of the Arkansas border (Summers et al.
1995), Dalea gattingeri was discovered by Logan in Fulton County, Arkan-
sas, in June, 1997 (Logan 97-20 [UAM, UARK}) and subsequently recorded
in the Arkansas Natural Heritage Commission data base from six additional
localities in that county, all dolomite glades (Logan 98-34 [UAM]). The
species resembles the widespread D. purpurea, differing in characters of the
inflorescence and bracts.

—_

Rhynchosia minima (L.) DC. This weedy vine was erroneously attributed
to the Arkansas flora by Grear (1978) based on a specimen from Aransas
(not “Arkansas”) National Wildlife Refuge in Aransas County, Texas (Smith
1988). Collections from the shores of the Mississippi River in Chicot County
here confirm its presence in the state. Thomas, Sundell, & Amason 142,827
(NLU).

Trifolium striatum L. Arkansas is one of a few southeastern states where
knotted clover, a European native, has had some success as an escape from
cultivation (Isely 1990). We record here four new localities from three counties,
nearly doubling its known presence in the state: Pulaski Co.: Thomas &

SUNDELL ET AL., Vascular plants from Arkansas 881

Amason 148,622 (NLU). Saline Co.: Thomas & Amason 148,835, 148,850
(NLU). Union Co.: Thomas & Amason 149,139 (NLU).

Vicia hirsuta (L.) S.F. Gray is a European native introduced to ruderal sites
over much of the United States (Isely 1990). The species has two seeds per
fruit and is often mistaken for the more common Y. ¢etrasperma, with four
seeds per fruit, and overlooked. In Arkansas, it has been documented only
from Washington County in the northwest. We add two southern counties
on the Louisiana border: Lafayette Co.: Thomas, Sundell, & Amason 156,549
(NLU). Union Co.: Thomas & Slaughter 104,337 (NLU, UAM).

GENTIANACEAE

Centaurium texense (Griseb.) Fern. Previous records of Texas centaury in
Arkansas were limited to a few counties in the Ozark Mountains, however
a recent collection from a blackland prairie site in Hempstead County in
southwest Arkansas (Swndell 12,368 [UAM}) represents an important (though
not unexpected) range extension within the state. Several species of calcar-
eous soils exhibit a similar pattern in Arkansas, for example, Juniperus ashe1
Buchholz and Penstemon cobaea Nutt.

IRIDACEAE

Sisyrinchium exile Bickn. Based on Hornberger’s (1987) merging of Szsyrinchinm
exile with S. rosulatum, Smith (1988) excluded the former taxon from the
Arkansas flora. Hornberger developed her concepts of Szsyrinchium species
in the southeastern United States “mostly from herbarium materials” (pers.
comm.). However, differences in stature and flower color between those taxa
that are immediately apparent in the field are obscure in dried specimens.
We recommend reinstatement of S. exé/e based on the following voucher
specimens: Ashley Co.: Thomas 92,053 (NLU, UAM). Bradley Co.: (where
plants of S. ex//e were mixed with the larger and more common S. roswlatum):

Sundell & Amason 11,737 (UAM). Union Co.: Thomas 133,595 (NLU).

LAMIACEAE

Clinopodium gracile (Benth.) Kuntze. Thieret reported C/inopodium grac-
ile as new to the United States in 1964 from a Louisiana collection. The
species has subsequently spread through much of Louisiana and into south-
ern Arkansas. Union Co.: Thomas & Amason 143,743 (NLU).

Scutellaria racemosa Pers. This recently introduced South American na-
tive is known from scattered localities on the Gulf Coastal Plain (Godfrey
& Wooten 1981). Originally picked up in Arkansas in 1992 by Marie Locke
during her work on the flora of Jefferson County (Smith, pers. comm.), its
presence in Arkansas is documented here by three collections, all from gar-

882 Sipa 18(3

Ww

dens, where the plants were likely contaminants in horticultural material
imported from further south. Drew Co.: Swndell 11,601 (UAM). Jefferson
Co.: E. & M. Sundell 12,055 (UAM). Union Co.: Thomas & Amason 155,860

(NLU). Time will tell whether this alien skullcap deserves resident status.

LILIACEAE

Erythronium albidum Nutt. Like bloodroot and toothwort, white trout
lily in Arkansas is acommon species of the Ozark and Ouachita highlands,
recorded by Smith (1988) for only two counties on the West Gulf Coastal
Plain. We here confirm his report for Arkansas County and add three sta-
tions from two other counties, all collections from richly wooded creek bottoms.
Arkansas Co.: Butcher s.n. (JAM). Cleveland Co.: Baker 20 (UAM). Drew
Co.: McDougald & Lincoln 15, Barbee, Lamb, & Pagan 36 (UAM).

LYTHRACEAE

Cuphea carthagenensis (Jacq.) J.F. Macb. was first collected in Arkansas
in Ashley County by Thomas in 1985 and recorded by Hooks (1986). Like
a number of others native to the Gulf Coastal Plain, the species is not unex-

2

pected in Arkansas’ southern counties. However, blooming from mid-summer
to fall, when field activities are typically less intense, it is probably
underrepresented in herbarium collections. We report it from a wet site in
Lafayette County in southwest Arkansas (Swndell, Thomas, & Amason 11,852
[UAM})

Didiplis diandra (Nutt. ex DC.) Wood. Water-purslane was collected at
Felsenthal National Wildlife Refuge in Ashley County (Swndell & McDonald
7623, 7630 [UAM}), where it grew both on mud and submerged in tem-
porarily flooded habitat. It was reported for the state (as Pep/is diandra Nutt.)
by Branner and Coville in 1891 but, to our knowledge, has never been
documented.

MAGNOLIACEAE

Magnolia macrophylla Michx. According to Tucker (1976), big leaf magnolia
grew as a native plant in Arkansas only in Clay County on Crowley’s Ridge
in the northeast corner of the state, where a single grove had been reduced
by natural hazards and local gardeners to a few small trees. Tucker predicted
that the species would soon be extirpated at the site; two trees remained in

1981 (Figlar 1981); a survey of the site in 1995 failed to relocate the spe-

cies (Meyer 1997). In 1994, Stuckey discovered a single tree of Magnolia
macrophylla growing on a heavily wooded slope above a stream, in the vi-
cinity of Mandeville near Texarkana in Miller County, in the southwest corner
of Arkansas (Stckey s.r. [UAM, UARK}). Wild populations of big leaf magnolia

in northern Louisiana are the most likely seed source.

SUNDELL ET AL., Vascular plants from Arkansas 883

MORACEAE

Fatoua villosa (Thunb.) Nakai. Sundell reported this herbaceous Asiatic
weed as new to Arkansas in 1986 based on a Drew County specimen. The
species has persisted in southern Arkansas (our most recent collection was
made in 1996), however, to our knowledge, it remains restricted to gar-
dens and nurseries, growing vigorously to heights of three feet. Pulaski
Co.: Sundell 10,440, E. & M. Sundell 12,049 (UAM). Union Co.: Sundell &
Amason 7,461 (UAM), Thomas & Amason 107,891, 111,269 (NLU).

PAPAVERACEAE

Sanguinaria canadensis L. Bloodroot is documented from almost every
county in the Ozark and Ouachita highlands of Arkansas (Smith 1988) but
is unknown from most of the southeastern half of the state. The phenom-
enon of more northern, highland species pioneering on the Coastal Plain
was discussed by Kral (1966), who noted that their random distribution on
some but not all richly wooded stream terraces in north Louisiana could
best be explained by impediments to dispersal rather than environmental
limitations. We record bloodroot from the mesic hardwood terrace of Hun-
ger Run Creek in Drew County where it grew with two other highland
associates, Cardamine concatenata (Michx.) Schwartz and Viola pubescens Att.:

Sundell & Pagan 10,152 (UAM).
POACEAE

Eriochloa acuminata (J. Presl) Kunth var. acuminata is known in Arkansas
at present from Conway and Mississippi Counties in the north central and
northeastern parts of the state (Smith 1988). Recent records from five addi-
tional counties suggest that the species is probably undercollected. Desha
Co.: Thomas, Sundell, & Amason 145,406 (NLU). Lafayette Co.: Thomas,
Sundell, & Amason 150,961 (NLU). Lee Co.: Thomas 134,402 (NLU). Miller
Co.: Thomas, Sundell, & Amason 151,107 (NLU). St. Francis Co.: Thomas
131,382 (NLU, UAM).

Rottboellia cochinchinensis (Lour.) Clayton. This noxious Asiatic grass
has become naturalized in the West Indies, Florida and Louisiana (Allen
1992). It appeared previously in Arkansas in Ashley County (Smith 1988)
and was rediscovered as a weed in a soybean field in 1996 in Arkansas County
by Brad Koen of the Arkansas Cooperative Extension Service. Koen s.n. (NLU,
UAM, UARK)

Setaria pumila (Poir.) Roem. & Schult. This distinctive, narrow-spiked,
European bristle grass was first reported for North America (as S. pallide-
fusca {Schum.} Stapf & Hubb.) from a Baton Rouge, Louisiana collection
by Thieret and Allen in 1974. It is currently known in that state from twenty

884 SIDA 183)

parishes (Thomas & Allen 1993). In addition, a collection at UAM docu-
ments its presence in Alabama. Setaria pumila has been found in weedy habitats
in three southern Arkansas counties. Ashley Co.: (1997): Sundell & Yezser
12,108 (NLU, UAM, UARK). Bradley Co.: (1993): Thomas 137,493 (NLU,
UAM). Calhoun Co.: (1998): Thomas & Amason 157,175 (NLU, UAM).

PORTULACACEAE

Portulaca umbraticola Kunth. Orzell and Bridges (1987) reported this
distinctive purslane from Monroe County in eastern Arkansas, and we here
note its occurrence at a second locality, in Miller County, in the southwest-
ern corner of the state. Thomas, Sundell, & Amason 151,119 (NLU).

PRIMULACEAE

Lysimachia japonica Thunb. was reported for Louisiana by Thomas and
Allen in 1982. A 1997 collection from Union County documents its pres-
ence in southern Arkansas. Thomas & Amason 155,858 (NLU).

PSILOTACEAE

Psilotum nudum (L.) Beauv. Whisk-fern has been reported by Peck and
Taylor (1995) for three counties in southwest Arkansas: Clark, Lafayette,
and Union. Thomas recently discovered the species at Grassy Lake in Hempstead
County (Thomas, Amason, Stuckey, et al. 155,572 {NLU]), where seventeen
plants were counted around the base of a large baldcypress tree. The species
is native in north Louisiana; in the older part of North Monroe, for example,
plants are common behind shrubbery that has not been replaced for several
decades. All such plants are less than six inches tall and do not get bigger:
they are the native diploid of Florida and the Gulf Coast, rather than the
much larger greenhouse tetraploid. Based on size, Thomas's Hempstead County
plants were diploids and more likely to be natives at the northern edge of
their range than waifs.

PTERIDACEAE

Pteris multifida Poir. in Lam. et al. Spider brake was first reported for Arkansas
in 1941 from Hot Springs National Park in Garland County, where it has
persisted to the present (Taylor 1984). We record a second locality for this
naturalized fern. Union Co.: Thomas & Amason 144,850 (NLU).

RANUNCULACEAE

Trautvetteria caroliniensis (Walt.) Vail. was discovered in Arkansas (on

Brady Mountain in the Ouachita Highlands, Garland County) in the late
1980's by John Pelton of the Arkansas Native Plant Society. Pelton com-

municated the discovery to Vernon Bates and Burt Pittman, who were then

conducting a floristic inventory of the Ouachita National Forest. Specimens

SUNDELL ET AL., Vascular plants from Arkansas 885

collected at that time have never been cited in publication as documenta-
tion of the population (Bates & Pittman, pers. comm.), however, it is likely
that the single Arkansas population of Trautvetteria mapped in Flora of North
America (Parfitt 1997) is the same. The monotypic genus is disjunct in North
America and eastern Asia, with North American populations of the eastern
and western forests recognized (rather arbitrarily according to Parfitt) at
the varietal level. Variety caroliniensis of the eastern United States was pre-
viously known to occur west of the Mississippi River only at several sta-
tions in Shannon County, Missouri (Steyermark 1963). Arkansas plants were
growing in abundance ona richly wooded seepage slope with Smilax lauvifolia,
Viburnum nudum, Osmunda cinnamomea, and O. regalis under a closed canopy

of mixed hardwoods and pine. Swndell 10,473 (UAM, UARK), 10,585 (UAM).

ROSACEAE

Pyrus calleryana Dene. When Sundell reported callery pear in 1986 as new
to Arkansas from Drew and Ashley Counties, the species was already wide-
spread on the state’s Coastal Plain as an escape from cultivation to early
successional habitats. The stout, thorny trees have proven to be a nuisance
to pine reforestation practices. We here document its status as a wild plant
in six additional counties, including two upland sites in the Ouachita Mountains.
Calhoun Co.: Sundell, Amason, & Etheridge 7,888 (UAM). Cleveland Co.:
Lunsford 12 (UAM). Hempstead Co.: Thomas, Sundell, & Amason 139,840
(NLU). Garland Co.: Swnde/l 10,650 (UAM, UARK). Miller Co.: Thomas,
Sundell, & Amason 151,152 (NLU). Montgomery Co.: E. & M. Sundell 12,357
(UAM). In addition, specimens at HSU document callery pear as adventive
in Clark County (Dan Marsh, pers. comm.).

SOLANACEAE

Datura ferox L. (D. guercifolia Kunth). This previously unreported species
of thorn-apple appeared in a soybean field in Clay County. It is a south-
western species (Correll & Johnston 1970) probably brought into our area
with agricultural seed and not likely to persist. (Thomas and McCoy re-
ported it from East Carroll Parish, Louisiana, in 1982.) Appreciation is expressed
to Andy Vangilder and John Boyd of the Arkansas Cooperative Extension
Service for sending material (Vanegz/der s.n.) to UAM.

VALERIANACEAE

Valerianella locusta (L.) Latterade was reported for Arkansas by Demaree
(1943) and Johnson (1971) but excluded by Smith (1988), who suggested
that Arkansas reports were probably based on cultivated material. It is re-
instated in the Arkansas flora with collections from a roadside in Sharp County,
where plants grew in abundance on mounds of soil (Swndell & Amason 12,074

[UAM, UARK}), and a graveyard in Howard County (Lawson 1507 [NLU]).

—

886 SIDA 18(3)

VIOLACEAE

Hybanthus concolor (T.F. Forst.) Spreng. In Arkansas, green violet is well-
documented from the Ouachita and Ozark highlands and from Crowley’s
Ridge. Collections from the wooded shores of Grassy Lake in Hempstead
C oe) mark its presence on the Gulf Coastal Plain. Swndell, Amason, & Stuckey
10,874 (UAM), Thomas & Amason 139,733, 155,576 (NLU)

ACKNOWLEDGMENTS
The authors express their appreciation to Isabel Bacon of the School of
Arts and Humanities, University of Arkansas at Monticello, for translating
the English abstract into Spanish.
REFERENCES

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Eunice, Louisiana.

BrANNER, J.C. and F.V. Covitie. 1891. A list of the plants of Arkansas. Ann. Rep. Geol.
Survey ee for 1888. 4:155-242.

Correitt, D.S. and M.C. JouNston. 1970. Manual of the vascular plants of Texas. Texas

Research See ees Renner.
Darwin, S.P., E. SUNDE es and A.S. BRADBURN. 1981. Noteworthy vascular plants from
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Demaree, D. 1943. A son of the vascular plants of Arkansas. Taxodium 1:1—88.
FIGLAR, R.B. 1981. A last stand in Arkansas. J. Amer. Magnolia Soc. 17(1):17—20.
GLEASON, H.A. and a: Cronouist. LODL. ee of the vascular plants of northeastern
United States and adjacent Canada, ed. e New York Botanical Garden, Bronx.
Goprriy, R.K. and J.W. Wooren. 1979. i and wetland plants of the southeastern
U biked ae - Monocots The Univ. of Georgia Press, Athens.
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ae

Dicotyledons. The Univ. of Georgia Press, Athens.

Grear, J.W. 1978. A revision of the New World species of RAynchosia (Leguminosae-Faboideae).
Mem. NewYork Bot. Gard. 31:1—168.

HoimGren, P.K., N.H. HoimmGren, and L.C. Barnerr. 1990. Index herbariorum. Part I:
The herbaria of the world, ed. 8. New York Botanical Garden, Bronx

Hooks, S.L. 1986. A Sree survey of the vascular flora of Ashley C ounty, Arkansas.
M.S. Thesis, Northeast La. Univ., Monroe.

Meier RGER, K.L. 1987. seat of the genus Sisyrinchinm oo in the south-
eastern United States. Ph.D. . Univ. of Arkansas, Fayettev

Isety, D. 1990. Vascular f
The University . North Carolina Press, Chapel Hill.

Jounson, M.I. 1971. A survey of the vascular flora of Poinsett County, Arkansas. Proc.
Arkansas Acad. Sc 25:42-52

Kartresz, J.T. 1994. A sy sees sce checklist of the vascular flora of the United States,
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Kral, 1966. Observations on the lop of the southeastern U aiid States with special
a

a rence to northern Louisiana. Sida 2:395—40

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ae YER, EG. 1997. Magnoliaceae. In: Flora of North America Editorial Committee, eds.
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at and Oxford. Vol. 3, pp. 3-10.

OrzeLt, S.L. and E.L. Brinces. 1987. Further additions and noteworthy collections in the
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PareitT, B.D. 1997. Trautvetteria. In: Flora of North America Editorial Committee, eds.

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PECK - H. and W.C. Tayzor. 1995. Checklis and distribution of Arkansas pteridophytes.
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888 Sipa 18)
BOOK REVIEW

FERNANDO SANTOS-GRANERO and FREDERICA BARCLAY (translated by Elisabeth
King). Selva Central. History, Economy and Land Use in Peru-
vian Amazonia. (ISBN 1-56098-761-8, hbk). Smithsonian Institu-
tion Press, 470 LEnfant Plaza, Suite 7100, Washington, DC 20560.
202-287-3738, ext. 343. 351 Pp.

This book is a translation, originally published in Spanish as, Ordenes y desordenes en la
Selva Central, in 1995. Many thanks should go to the Smithsonian Tropical Research Insti-
tuce, and especially Ira Rubinoff, for encouraging its translation

he book covers the Selva Central region of Peru, also called the “central montafia, com-
posed of portions from the western portions of: Oxapampa Province, in the Department of
Pasco, and Chanchamayo and Satipo Provinces in the Department of Junin. This is one of
the ewo most biologically interesting parts of the country because of its myriad of habitat
jalca” on sandstone containing Guayana Highland floristic ele-

“

types, from high altitude
ments, to middle elevation “ceja de selva,” largely comprised of cloud and montane forest
with many endemic species. The book is largely a socio-economic history of the region,
but it also contains extremely valuable information on the ecological characteristics, the
effects of agropastoral and logging activities, and other aspects related to land tenure, in-
cluding the unique “minifundios.”

The book is divided in three parts: genesis of a regional space, the colonist economy, and
indigenous integration. While this kind of book may not a normal sae is for the
everyday monogr apher or one anyone concerned with long-term conservation of
any tropical area is well-served by reading this. I have pele ae to establish long-term
studies, it 1s critical to see the evolution of the local populations, (be they indig-
enous or otherwise) so that our in-country counterparts might better translate our con-
cerns for basic science into information for the applied world of the local residents. The
most important part for me was the second one, especially chapters four, five and six, where
the land tenure systems, patterns and intensity of land use, and finally, economic articula-

n and environmental degradation themes are discussed, respectively. Having worked in
Peru since 1990, reading of this book gave me a new appreciation for the cela s of
managing the biotic resources of the region. While several portions of the “conclusions”
chapter are now outdated, an support a particular political point of view, the three chap-
ters of the second part certainly make this book a worthwhile investment for any botanist
working in the Andes, and especially, in Amazonia. This book is not only appropriate for
sotanical libraries, but also for ethnobotanists, and for anyone assisting in

—

acquisition by
the establishment of Integrated Conservation and Development areas.—John J. Pipoly HL

Stipa 18(3): 888. 1999

NOTEWORTHY RECORDS OF MISSISSIPPI
VASCULAR PLANTS

BRUCE A. SORRIE

160 West Rhode Island Ave.
Southern Pines, NC 28387, U.S.A.

STEVE W. LEONARD

CSTS-ENV, Bldg. 6678
Camp Shelby, MS 39407-5500, U.S.A.

ABSTRACT

Herbarium ee and field exploration, primarily in the Longleaf Pine and Coastal Pine
Meadows regions, have yielded the following new Species to a Mississippi: Chamate vista deeringiana,
Cladinm mariscoides, eae sas fusiforme, Dryopteris ludovi ocharis elongata, E. melanocarpa,
. robbinsit, Lsoetes loutstanensis, Lobelia boykinii, Mitreola cael ee decurren

. globularis var. pinetorum, R. harperi, a ae SS Sagittaria isoetiformts, ee
ees Sensi stricto, oe brevilabris var. floridana, Utricularia olivacea. The following
are previously known from the state, but are ee oll indy or represent significant rang
eee within Mississippi: Agrimonta incisa, Aristida condensata, A. simpliciflora, A. tuberculosa,
Burmannia biflora, Cal mssliafloras, Coelorachis cylindrica, enaamn 5 ere. ale Elyoneurus
tripsacoides, Gordonia lasianthus, Lsoetes melanopoda, Juncus gymnocarpus, Marshallia trinervia,
M eas laxum, Parnassia grandifolia, Poly gala leptostachys, Potamogeton am Rhynchospora

tis, R. fascicularis var. distans, R. nitens, R. tracyi, Ruellia pedunculata ssp. pinetorum, Sorghastrum
peas Stylisma aquatica, 8. pickeringii, Tridens carolinianus.

RESUMEN

Los estudios de herbario y ex sais de campo en las zonas de pino palustre y de
y

prados costeros con pinos, han nuevas para Mississippi: hoi

ah ae Cladium mariscoides, ee fs aoe Dryopteris dia

's elongata,

nelanocarpa, E. robbinsii, Isoetes lowisianensis, Lapalabeing Mitreola angustifolia, Rh Ue
dros R. ee var. RT - pain Rhy membre ed a sae isoetiformi
var. floridana, Ut . Las siguientes

se conocian previamente del. estado, Lay se colectan raramente o representan una ampliacién
significativa de su areal er Mississi ania incisa, Aristida condensata, A. simpliciflora,
A. tuberculosa, Burmannia Liflora, Ci ees Coelorachis cylindrica, Dichanthelium
erectifolinm, Elyoneurus trips.tcotdes, is ordonta lasianthus, lsoetes melanopoda, Juncus gymnocarpus,

ae}

Marshallia trinervia, Myriophyllum laxum vassia grandifolta ieee Ds Potamogeton
epihydrus, Rinna ae 1, R. ra ris var. distans, R. nitens, R. tracyt, Ruellia peduncilat.
ssp. pinetorum, Sorghastrum sepia Stylisma aquatica, 8. pickeringii, Tridens carolini

INTRODUCTION

Field work was conducted during 1995-98, primarily in a ten county
area of southeastern Mississippi (Forrest, George, Greene, Hancock, Harrison,

Sipa 18(3): 889-908. 1999

890 SIDA 1834)

Jackson, Lamar, Pearl River, Perry, and Stone counties). From 1995—96 work
was carried out by Bruce A. Sorrie (BAS) under a contract from the United
States Fish and Wildlife Service, Endangered Species, to survey seepage bogs
and wet savanna habitats of southern Alabama and Mississippi for selected
Federal Species of Concern (formerly C2 candidates). In 1997, a rare spe-
cies inventory was conducted by BAS on the 22,000+ acre University of
Mississippi Forest Lands in George, Jackson, and Stone Counties, under a
contract from The Nature Conservancy, Mississippi Field Office. From 1997-
98 work was carried out by BAS under a contract from The Nature Conser-
vancy, Southeast Region Office, to survey remaining high quality longleaf
pine (Pinus palustris P. Miller) communities.

In 1993 the Mississippi Military Department signed a memorandum of
agreement with the Mississippi Department of Wildlife, Fisheries, and Parks
for a biological inventory of Camp Shelby Training Site (CSTS). Camp Shelby
is located principally in Forrest and Perry Counties, and covers 134,000
acres, mostly leased from the DeSoto National Forest. The Mississippi Museum
of Natural Science and the state’s Natural Heritage Program hired contrac-
tors and coordinated field work at CSTS. From 1995-98 Steve W. Leonard
(SWL) was field botanist on the inventory team.

Southeastern Mississippi lies within the Longleaf Pine and Coastal Pine
Meadows Physiographic Regions, as defined by Lowe (1921). Although

em

supporting a wide variety of habitat types, varying from xeric maritime dune
scrub to salt marshes to mixed mesophytic ravine forests, the area is most
notable for its extensive pine forests, pine savannas, and seepage bogs. Southward,
and in wet sites northward, slash pine (Pinus e//iottii Engelmann) is domi-
nant; while northward, and in mesic to dry sites southward, longleaf pine
is dominant. Mesic and moist sites usually support a mixture of the two.
Beneath the pines is a fire-adapted, species-rich herbaceous layer dominated
by grasses, composites, and legumes. Wet sites usually also support a dense,
ericaceous shrub layer. North of the six southernmost counties, loblolly pine

(Pinus taeda L.), shortleaf pine (Pinus echinata P. Miller), and to a lesser ex-
tent spruce pine (Prnus glabra Walter) become important components as
the topography becomes more rolling and dissectec

The dramatic loss of the longleaf pine ecosystem over 97% of its former
range has been documented by Ware et al. (1993) and Frost (1993). Al-
though similar losses occurred in Mississippi, much of the southeastern portion
of the state remains ina relatively natural condition. This is due to the presence
of large public lands and the implementation of recurring fire management.
The De Soto National Forest (which includes Camp Shelby within its bor-
ders), the Mississippi Sandhill Crane National Wildlife Refuge, and the University
of Mississippi Forest Lands together form one of the largest and most eco-

—
—

logically significant blocks of longleaf pine ecosystem remaining (222,000

SORRIE AND LEONARD, Mississippi vascular plant records 891

ha). Many of the taxa reported in this paper inhabit fire-maintained com-
munities within these properties.

Knowledge of the flora of Mississippi is less well known than most other
southeastern states (Bryson & Carter 1994). Published or draft atlases of
vascular flora exist for Arkansas, Florida, Georgia, Louisiana, South Caro-
lina, and Tennessee. Major revisions of state floras are well advanced in the
Carolinas and Virginia (combined) and Texas. In Mississippi, and also in
Alabama, a paucity of historical collecting coupled with a diverse and
biogeographically complex flora has hindered the publication of modern
floristic manuals. Lowe (1921), Small (1933), and Radford, Ahles, and Bell
(1968) remain as basic references for Mississippi plants. The Flora of Mis-
sissippi Project was begun in the 1960s (Jones 1974) and has resulted in
treatments of a number of families and genera [see references cited by Stevens
(1991) and Bryson et al. (1996)}. Numerous county floras have added greatly
to our knowledge of plant distribution (see Stevens 1991 for most titles).
Currently, the PLANTS database provides a comprehensive list of taxa docu-
mented from the state (USDA, NRCS 1995). This database admits species
based on vouchered specimens, monographic treatments, and literature reports.

In this paper, plants are listed alphabetically by genus and species. No-
menclature follows Kartesz (1994) unless noted otherwise. Herbarium ab-
breviations follow Holmgren et al. (1990), except for bas, ctb, MMNS, and
swl (pers. herb. Bruce A. Sorrie, pers. herb. Charles T. Bryson, herbarium
of the Mississippi Museum of Natural Science in Jackson, and pers. herb.
Steve W. Leonard). Historical specimen data was derived from the Missis-
sippi Natural Heritage Program database (MSNHP) and from rare species
files developed by BAS. Specimen label data was transcribed exactly as the
collector wrote it, except that lists of associated species have been omitted.
For some taxa, where we have documented many current populations, only
a few representative collections are cited. The abbreviation FS stands for

Forest Service Road.

NOTEWORTHY COLLECTIONS

Agrimonia incisa Torrey & Gray (Rosaceae). Previous Mississippi collec-
tions are from Forrest, George, Harrison, and Simpson Counties (MSNHP);
to these we add Perry and Stone Counties. Once considered a candidate for
listing by the US Fish and Wildlife Service, incised groove-bur has a spotty
distribution from South Carolina to central Florida to eastern Texas. Camp
Shelby, straddling Forrest and Perry Counties, supports ten discreet occur-
rences. Habitats vary from mesic longleaf pine woodland to entrances to
gopher tortoise burrows in dry pine-oak woodland, pine plantations, and
cemeteries; but it is most abundant on roadside embankments. MacRoberts
and MacRoberts (1997) cite similar habitats for Agrimonia incisa in Texas

892 SIDA 18Q)

and suggest that fire suppression and subsequent habitat alteration have
caused a widespread decline in this species.

Voucher specimens: Perry Co.: Camp Shelby, frequent bee MS hwy. 29 in gr roe ad
shoulder, 5.4 mi S of US 98 at New Augusta, 12 Jun 1996, 8. W. Leonard 9543 (FSU

NCU); Camp Shelby, Training Area T-51, 28 Dec 1995, S. W. Leonard 9363 een Stone
Co.: Univ. Miss. Forest Lands, dry-mesic oa ate upland with hardwood understory, N of
Wire Road and W of Cablebridge Road, 22 Aug 1997, B.A. Sorrie 9455 with J. Moore (bas

Ww

Aristida condensata Chapman (Poaceae). Lowe (1921) does not include
this species of dry to xeric sandhills, but PLANTS does. Mississippi popu-
lations mark the western edge the species’ range.
nigel specimens: Greene Co.: Leaf River at hwy. 98, 2 Nov 1969, K.E. Rogers 2510
(NCU, B). Harrison Co.: longleaf pine-oak-saw palmetto ridge on FS 415-A, De Soto
Seer ae frequent, 13 Aug 1997, B.A. Sorrie 9393 (MMNS, NCU). Perry Co.: infrequent
in cut-over longleaf pine-oak scrub behind Mars Hill Church, Camp Shelby, 11 Jan 1996,
5. W. Leonard 9376 (MMNS), same place, 9 Oct 1996, 8. W. Leonard 9660 (NCU). Wayne
: common in xeric longleaf pine-turkey oak sandhills by Brewer Town Road, De Soto
al Forest, 26 Sep 1997, B.A. Sorrie 9552 with M. Pyne (GH, NCU
Aristida simpliciflora Chapman (Poaceae). This rare but easily overlooked
grass is known from less than 25 counties range-wide (central Florida to
southern Mississippi). Rogers (1977) collected it from the Ragland Hills in
Forrest County; the following are the remaining collections known to us.
Voucher specimens: Pearl River Co.: McNeill, 18 Nov 1929, H.R. Reed s.n. (US); four
mi S of Poplarville on US 11 at jet. of MS 26, base of longleaf pine slope, local, 30 Sep
1988, S. rie 30289 (IBE, VBD). Perry Co.: Camp Shelby, fairly abundant in aiechen
plant seep g a secondary drainage into Flat Creek, 3.7 mi S of FS 385, 23 Oct 1996,
SW aes ee (MMNS).
Aristida tuberculosa Nuttall (Poaceae). Lowe (1921) does not include this
species of semi-stable dunes and xeric sandhills, but PLANTS does. Aristida
tuberculosa has a unique distribution pattern: upper Mississippi River and
immediate tributaries; about the head of lake Michigan; coastal New Hampshire
to Delmarva Peninsula; fall-line sandhills of the Carolinas and Georgia; plus
scattered locations on the coastal plain from South Carolina to Florida and
Louisiana. Some of these latter may represent adventive populations in dis-
turbed soils and fallow fields.

—

Voucher specimens: Forrest Co.: James Street, E of Baptist Church, lake at old gravel
and sand pit, in sandy open woods, 1971, K.E. Rogers 7028 (VDB), 7030 (NCU). Kemper

».: 5 mi S of DeKalb, sandhill with Quercus incana, Q. margarettiae, a few Pinus palustris,
13 Sep 1971, 8. McDaniel 15560 IBE, VDB). Jackson Co.: Horn Island, 5 Sep 1891,
S.M. Tracy 1579 (US). Wayne Co.: common in xeric, deep white sands of hills along Brewer

Town Road, De Soto National ee ae Pinus palustris, Quercus laevis, Aristida condensata,
ce megalocarpa, 26 Sep 1997, B.A. Sorrie 9553 and M. Pyne (GH, MMNS,

SORRIE AND LEONARD, Mississippi vascular plant records 893

Burmannia biflora L. (Burmanniaceae). This endemic to the Atlantic and
Gulf Coastal Plains has rarely been collected in Mississippi. Jones (1976)
lists four counties, but the specimen that he saw from Jackson County ac-
tually is Apteria aphylla {Channell 1661 (MISSA)}. Jones also lists Pearl River

o., but we have not yet located a voucher. The primary habitat in Missis-
sippi is seasonally ponded depressions and, occasionally, seepage bogs.

Voucher eens Hancock Co.: Bay St. Louis, Nov 1905, A. A/fison 1425 (MISS,
photocopy at IBE). Harrison Co.: soe in sphagnum tussocks in Nyssa-Sphagnum swamp
along ae aa River, 3 mi N of Seymour, 20 Aug 1952, J.D. Ray, Jr. ci (FSU,
MISSA, NCU). Jackson Co.: Oren ca 7 Sep 1889, FS. Earle s.n. (DUKE).
Calopogon multiflorus Lindley (Orchidaceae). Lowe and PLANTS do not
list C. multiflorus for Mississippi, but Luer (1972) maps it there. This essen-
tially Floridian orchid has rarely been collected outside of that state.

Voucher specimens: Hancock Co.: three mi south of Necaise, 5 Apr 1967, eS a Jones, Jr.
11489 (MISS). Jackson Co.: Ocean Springs, 4 Apr 1888, ES. Earle s.n. (AMES, ie Ocean
Springs, Apr 1893, J. Skehan 206 6 (BRIT): two mi W of Alabama state line, au
R.D. Suttkus 66-2-11 (NO); circa | mi W of MS/AL line between I-10 and US 90 on eae
Creek Road, recently burned savanna, rare, 20 Apr 1984, C. Norquist 1774 (IBE); ca. 6.3
mi E of Moss Point, plants scattered in pine savanna, 15 Apr 1988, M.W. Morris 3076 et al
(IBE)

Chamaecrista deeringiana (Small & Pennell) Macbride (Fabaceae). Re-
ported herewith new to Mississippi. The following establishes a new west-
ern range limit for this essentially Floridian species.

Voucher specimen: Harrison Co.: Deer Island, W end, live oak woods over sand or shell,

29 Aug 1977, M.B. Brooks 432 with 8. McDaniel (IBE).
Cladium mariscoides (Muhl.) Torrey (Cyperaceae). Reported herewith new
to Mississippi. This species was not included for Mississippi by Bridges et
al. (1993). This species occurs primarily within the southern portion of the
glaciated region of eastern North America, with scattered occurrences in
most southeastern states.

Voucher specimens: Jackson Co.: swamp ponds, Ocean Springs, 19 Aug 1953, D. Demaree

33953 (GH); pine meadows between hwy. and RR, about halfway between Pascagoula and
Fontainbleu {sic], 22 Aug 1962, R.M. Harper 4530 (GA, GH); abundant on boggy margin
of pond on N side of John Smith Road, just W of Huey Davis Road, 26 Sep 1996, B.A,
Sorrie 9066 with R.J. LeBlond (MMNS, NCU): abundant in open wet slash pine flatwoods
on E side of Trent Lott Airport, N of Sarracenia Road, 11 Nov 1997, B.A. Sorrie 9633
(IBE, NCU)
Coelorachis cylindrica (Michaux) Nash (Poaceae). Although there are numerous
records for Arkansas and Louisiana, this grass seems rare east of the Missis-
sippi River. PLANTS lists this grass for the state, but the following are the
only Mississippi specimens we are aware of.

894 Sipa 18(3)
Voucher specimens: Forrest Co.: 1/2 to 1 mi N of Petal, between N Railroad St. and
hwy. 11, several plants at this site, 23 Jul 1971, K.E. Rogers 6720 (NCU). Monroe Co.:
just N of Lowndes County, where US 45 crosses Buttahatchee River, on roadside bank, 30
Jun 1991, J. MacDonald 2771 (IBE).
Dichanthelium erectifolium (Nash) Gould & Clark (Poaceae). Lelong (1986)
reports this grass only from Jackson County, the western range limit of the
species. It is apparently rare, for we have not yet encountered a Mississippi
specimen in searches of over a dozen southeastern herbaria. Once learned,
Dichanthelinm erectifolium is easy to distinguish from other congeners, and
in Mississippi is restricted to seasonally ponded cypress depressions, usu-
ally acompanied by D. wrightianum, Polygala cymosa, Sabatia bartramit, [ex
myrtifolia, and Hypericum myrtifolinm.

Voucher specimens: Harrison Co.: abundant in seasonally ae etoressios in pine
Hatwoods off of FS 452, S of Wortham Road, De Soto National Forest, grass-sedge domi-
nated, with Nyssa biflora, ene wrightianum and Cyesnodille 5 aera 14 Aug
1997, B.A. Sorrie 9404 (G

side of Frank Snell Road, 7 7 route 613, abundant in pond cypress depressions in pine

flatwoods, 5 Jun 1996, B.A. of sn. (ba

3, NCU, MMNS). Jackson Co.: Cottonmouth Savanna, N

Dichanthelium fusiforme aie Harvill (Poaceae). Reported here-
with new co Mississippi. This combination was made by Harvill (1977),
but is not included in synonymy under D. aciculare by Kartesz (1994). This
plant’s taxonomic status has long been in question, and recent authors have
treated it as D. aciculare var. ramosum (Grisebach) Davidse (Davidse & Pohl
1992) or Panicum aciculare var. arenicoloides (Ashe) Beetle (Zuloaga et al, 1993).
Lelong (1986) synonymized Panicum fusiforme under a related species, P.
angustifolium Elliott. However, the very long (3.0 mm) fusiforme spikelets
are strikingly different from the blunt spikelets of D. aciculare and D, (Panicum)
angustifolium, and so we will use Harvill’s combination until a comprehen-
sive revision is undertaken. The following is the only specimen we have seen
for Mississipp1.

Voucher specimen: Jackson Co.: thickets near the coast, Ocean Springs, 1952,
D. Demaree 32226 (GH). The spoomicn, was originally determined to be
Panicum angustifolium, but spikelets are 3.0 mm long and fusiform.

Dryopteris ludoviciana (Kunze) Small (Dryopteridaceae). Reported herewith
new to Mississippi. This coastal plain endemic was not included for Missis-
sippi by Evans (1978) or by Montgomery and Wagner (1993).

Voucher specimens: Perry Co.: Camp Shelby, about 100 plants on W side of Denham
Creek, 2 air mi southwest of New Augusta, 14 Sep 1995, S.W. Leonard 9082 (MMNS).
Eleocharis elongata Chapman (Cyperaceae). Reported herewith new to
Mississippi. Although ranging from North Carolina to Florida and Texas
(and scattered in the neotropics), this species is not listed by Lowe (1921)
or PLANTS. This sedge usually inhabits seasonally to permanently ponded

SORRIE AND LEONARD, Mississippi vascular plant records 895

depressions. The rhizomes are unusually well developed in the Ocean Springs
specimen, and the habitat is also unusual, but the specimen appears to be
correctly identified, with slender spikes as wide as the culms and with red-
margined scales.

Voucher specimens: Hancock Co.: Stennis Space Center, NASA, 13 km SE of Picayune,
main canal near N-S roadway, 8 eek 992, ].W. Wooten s.n. (USM). Jackson Co.: 7 mi E of
Ocean Springs, local colony on moist slope near longleaf covered hill, 27 May 1962, S.
McDaniel 3231 (BE); Horn Island, eran in freshwater marsh shortly S of Park Service
pier, 3 Oct 1997, B.A. Sorrie s.n. with R.J. LeBlond (bas). Lamar Co.: Mossy Pond, 3 mi W
of Purvis, uncommon, with E. robbinsit, | Sep 1998, B.A. Sorrie 9932 with S.W. Leonard

U

Eleocharis melanocarpa Torrey (Cyperaceae). Reported herewith new to
Mississippi. This species is not listed by Lowe (1921) or PLANTS. Unlike
other Mississippi species, the tips of the culms of E. melanocarpa often arch
over and root in the moist or wet sandy substrate, thus forming a dense
tangle. This sedge ranges from Massachusetts to Florida and Mississippi,
disjunct to eastern Texas, southern Michigan, and northern Indiana.

Voucher specimen: Perry Co.: Camp Shelby, abundant in nearly dry gum pond behind
Mars Hill Church, 23 Apr 1995, J. MacDonald 8469 with A. Leidolf nee same place, 9
Oct 1996, S.W. Leonard 9656 with J. Moore and J. MacDonald (FSU, MMNS).
Eleocharis robbinsii Oakes (Cyperaceae). Reported herewith new to Mississippi.
Although known from the state for over twenty-five years, this is the first
published documentation. Mississippi marks the western range limit for
the species. Plants usually form dense beds in shallow gum ponds with fluctuating
water levels; two such ponds on Camp Shelby support populations. In con-
trast to E. elongata, the scales of E. robbinsiz are green, often with a translucent
scarious margin, and without any red coloring. Statements to the contrary in
Godfrey & Wooten (1979) were based on misidentified specimens. In addi-
tion, scale length in E. e/ongata is 3.5-4.5 mm, in E. robbinsit 5.5-7.0 mm.

Voucher specimens: Forrest Co.: hwy. 49 about 3-5 mi N of Stone Co. line, shallow
pond west of highway across fence, 30 Sep 1970, K.E. Rogers 4658-B (NCU); same pace
but actual distance is 1.2 mi N of county line, 19 Jun 1998, B.A. Sorrie 9780 (GH
MMNS); vicinity of Maxie, edge of boggy area, 5 Jul 1971, K.E. Rogers 6662-B nce

amar Co.: about 3 mi W of Purvis, abundant, 29 Aug 1973, K.E. Rogers 9225 (NCU);
same place [Mossy Pond], 1 Sep 1998, B.A. Sorrie 9931 with S.W. Leonard (GH, bas). Perr
Co.: Camp Shelby, very abundant around edge of gum pond behind Mars Hill Chao 20
Sep 1994, J. MacDonald 7728 with D. Wyrick IBE); same place, 11 Sep 1995, S.W. Leonard
9059 (FSU, MMNS).
Elyoneurus tripsacoides Humboldt & Bonpland ex Willd. (Poaceae). This
primarily neotropical species is not listed by Lowe (1921), but is included
in Hitchcock (1950) for Mississippi. That report may be based on a dupli-
cate of the following. In the United States it is also known from Florida
and southern Georgia.

896 Sipa 18(3)

Voucher specimen: Harrison Co.: Biloxi, 21 Aug 1898, S.M. Tracy 4594 (NCU).
Gordonia lasianthus (L.) Ellis (Theaceae). Lowe (1921) cites loblolly bay
only from Bay St. Louis in Hancock County. From then until 1994, only
12 sites were documented in George, Perry, and Stone Counties (MSNHP).
Extensive searches on Camp Shelby have revealed it to be far more com-
mon—25 occurrences have been located there. All of these populations consist
of small numbers of plants, primarily in seepage of toe slopes and along
streams where organic soils are present. Mississippi marks the species’ western
range limit.

Voucher specimens: ee Co.: bottomland at base of slope, E of dirt road in T2S RSW
Sec. 26, 17 Nov 1978 1. Rogers 46208 et al. (BE, MMNS). Perry Co.: Camp Shelby,
secacnar i in margin se pees forest on W side of Deep Creek, near S end of FS 332-A,
originally discovered by D. Wyrick, 8 Sep 1995, S.W. Leonard 9048 with J. MacDonald
(MMNS), /. Mac i ene io Stone Co.: about 1/4 mi S of McHenry, 6 Aug 1952,
R.L. Diener 375 (MISS

&

—

Isoetes louisianensis Thieret (Isoetaceae). Reported herewith new to Mississippi.
This quillwort was described from southeastern Louisiana by Landry and
Thieret (1973). Prior to 1996, I. /owisianensis was known from only eight
populations in Washington and St. Tammany Parishes and was listed as
endangered by the US Fish and Wildlife Service. In 1996 we independently
collected unknown quillworts in the De Soto NF and Camp Shelby, which
were verified as I. /owisianensis by W. Carl Taylor and Neil Luebke of the
Milwaukee Public Museum. Subsequently, an intensive search was conducted
throughout the De Soto NF by Forest Service personnel and SWL, and elsewhere
by SWL. To date, over SO populations have been found in ten Mississippi
counties. This species normally inhabits intermittent streams and stream
edges, scour channels, and floodplain depressions beneath a canopy of Owercus
laurifolia Michx., Nyssa biflora Walter, Acer rubrum L., and Cyrilla racemiflora
L. Soils are mineral in content. Plants initiate growth during winter months
and produce mature megaspores from April to July. Around mid-July, streams
dry up and plants senesce and disappear until rains resume in November.
In addition to the following, Louisiana quillwort has been vouchered by
SWL from Hancock, Harrison, Pearl River, and Stone Counties.

—

Voucher specimens: Forrest Co.: Camp Shelby, go ick in tributary of Chaney Creek,
27 Jan 1997, S.W. Leonard 9729 (NY). Greene Co requent in tributary of Waterfork
Branch, 8.6 mi E of Bothwell on N side of Bothwell ae de Road, 19 Jan 1997, S.W.
Leonard 9728 (NY). Jackson Co.: De So ‘local in braided streamhead of Bayou Billie,

side of Old Biloxi Road, 18 Jun 1 906, B. a Sorrie 8932 (GH, MIL). Jones Co.: ae Soto
on shallow still water of scour channels $ of FS 201, 0.5 mi SE of jct. with FS 201-F, 15
Feb 1997, 8S. W. Leonard 9743 (NY). Perry Co.: Camp Shelby, abundant in scour veeer
at edge of floodplain of tributary of Joes Creek, Be ee hardwood bottomland, 22 May
1996, S.W. Leonard 9511 (MMNS, NY). Wayne Co.: to NE, abundant in Okey Branch,
E of hwy. 63 and S of FS 207, 15 Feb 1997, S.W. Ee) O(NY).

SORRIE AND LEONARD, Mississippi vascular plant records 897

Isoetes melanopoda Gay & Durieu (Isoetaceae). Taylor et al. (1993) in-
clude Mississippi in the range, but Evans (1978) does not and we have seen
no earlier collections than those below. This quillwort grows in moist mixed
hardwoods and in ephemeral streams, generally north of the range of 1. /owisianensis.
Voucher specimens: Jasper Co.: Bienville National Forest, ephemeral stream S of FS
507, between FS 507-F and 507-C, 7 Apr 1997, : W. Leonard 9787 (NY). Scott Co.: Bienville
NE, occasional in ephemeral stream W of FS 543, 3.1 mi SE of jet. with FS 551, 7 Apr
1997, S.W. Leonard 9792 (NY). Smith Co.: oe NE, scarce in shallow depression of
floodplain, N side of FS 520, 0.4 mi E of Ichusa Creek and 2.1 mi E of county road 515, 7
Apr 1997, S.W. Leonard 9793 (NY). Wayne Co.: frequent in willow oak-sweetgum-winged
almetto forest on E side of US 45, S of jct. of Hayes-Nute Road, between Hiwannee

and abe: 14 Dec 1996, 8. W. Leonard 9718 (NY).
Juncus gymnocarpus Coville Juncaceae). This rush is not listed by Lowe
(1921) nor by Pullen et a/. (1968). Rogers (1973) is apparently the first to
report this rush from the state. The geographical range consists of three
disjunct areas: montane eastern Pennsylvania; montane North Carolina, South
Carolina, and Tennessee; and the coastal plain of panhandle Florida, south-
ern Alabama, and southern Mississippi. The long disjunction between the
Appalachian Mountains and the Gulf Coastal Plain suggests the possibil-
ity of taxonomically different entities, but R. Kral has seen specimens from
Camp Shelby and concurs with our identification (pers. comm. toSWL 1996).
To date sixteen occurrences have been documented from Camp Shelby, and
four from UMISS Forest Lands. In addition to the following, McDaniel (1987)

reports it from Lamar County.

Voucher specimens: Forrest Co.: hwy. 98 a few mi SE of junction with hwy. 49, numer-
ous at a of farm pond, 26 Oct 1972, K.E. Rogers 8841-E (MISSA); large colony in black
gum-sweetbay- Ae eee forest along Poplar Creek, Camp Shelby, 26 Sep 1996, S.W.
Leonard 964 9 (MMNS). Perry Co.: occasional along West Fork of Denham Creek, ‘e amp

Shelby, 18 Sep ee W. Leonard 9088 (MMNS); locally abundant on E side of Sweetwater
Creek, N of FS 305-F1, Camp Shelby, 16 Apr 1996, S.W. Leonard 9428 (NCU). Stone Co.:
wet seepage in mesic hardwood-loblolly pine ravine by Long Branch, N of Wire Road,
Univ. Miss. Forest Lands, 12 Jun 1997, B.A. Sorrie 9278 (GH, MMNS).

Lobelia boykinii Torrey & Gray ex A. DC. (Campanulaceae). Reported herewith
new to Mississippi, disjunct from Houston County, Alabama, and Okaloosa
County, Florida. This coastal plain endemic is most numerous in Georgia
and the Carolinas, with a few populations in New Jersey, Delaware, the Florida
panhandle, and Alabama.

Voucher specimens: Jackson Co.: Cottonmouth Savanna, N side of Frank Snell Road, E
of route 613, locally numerous in pond cypress depressions in slash pine flatwoods, 5 Jun
1996, 10 Nov 1997, and 19 Jun 1998, B.A. Sorrie 8871 (GH, NCU, MMNS), 9785 (FSU,
a uncommon in disturbed pond cypress depression at junction of Nut Bank Road and
ector to route 613, about 3.5 mi SSW of Big Point village, 2 Oct 1997, B.A. Sorrie
ee with R.J. LeBlond (bas).

898 Sipa 18(3)

Marshallia trinervia (Walter) Trelease (Asteraceae). This striking mem-
ber of the genus was brought to the attention of SWL by Dr. Sam Rosso of
the University of Southern Mississippi, who reported a station near the Lamar/
Marion County line. Despite knowledge of its presence in Mississippi for
nearly 150 years, M. trimervia has traditionally been considered rare in the
state. Lowe (1921) lists it only from Greene and Tishomingo Counties, but
Channell (1957) cites collections from Clarke, Forrest, Greene, Perry, and
Scott Counties. Watson and Estes (1990) add Covington, Lee, and Pearl
River Counties. At the University of North Carolina herbarium (NCU) there
are specimens from seven sites in Covington, Forrest, and Pearl River Counties.
Thus, it would seem that this species is not as rare in Mississippi as previ-
ously thought. Primarily a plant of creek banks, wooded slopes, and lime-
stone cliffs from central Alabama and western Georgia to central Tennes-
see, recent collections suggest that Marshallia trinervia has exploited moist
roadsides in the Gulf Coastal Plain. We here report stations not previously
documented.

Voucher Se Forrest Co.: ae 11 and 49 cloverleaf, cleared area, 2 Jun 1965,
S.B. Jones, Jr, 2935 (NCU); about 1/2 mi S of Hattiesburp, edge of low pine woods, 24 Aug

1969, K.E. Rogers 1350- c (NCU). Lamar Co.: moist roadside, 6.0 mi : of US 98 on W

side of MS hwy. 89 and 0.4 mi S of junction of WPA Road, N of Purvis, 22 Jun 1997, S.W.
relies 3 (MMNS). Pearl River Co.: 4.4 mi S of Derby on US 11, eee and ditches,
30 May 1976, D.E. Boufford 18652 et al (NCU), wet ditches, hwy. 11, 1 mi N of McNeill,
2 Jun ie C.F Reed 103625 (NCU); low area on roadside of hwy. 26, about 15 mi E of
Poplarville, 30 May 1976, G. Nesom s.n. (NCU) {this location probably is in Stone Co.].
Perry/Forrest Co.: Camp Shelby, edge of shrubs along small streams on E and W sides of
Poplar Creek, at county line, 22 May 1998, S.W. Leonard 9959 (MMNS).
Mitreola angustifolia (Torrey & Gray) J.B. Nelson (Loganiaceae). Reported
herewith new to Mississippi. Godfrey and Wooten (1981) credit this plant
only to southern Georgia and northern Florida. Mitreola angustifolia inhab-
its seasonally ponded depressions in pine flatwoods, often under a thin canopy
of pond cypress.

Voucher specimens: Jackson Co.: savanna circa 1 mile W of Alabama state line on S

side of US 90, 26 July 1983, C. Norguist 1291 (IBE). Scattered in drying out cypress pond
on E side of hwy. 63, 14.4 mi S of jet. of hwy. 26 in Lucedale, 4 Aug 1996, S.W. Leonard
9596 (MMNS, NCU); Cottonmouth Savanna, N side of Frank Snell Road, E of route 61
cypress-bordered depression in pine flatwoods, 1 Oct 1997, B.A. Sorrie and R.J. LeBlond
95835 (GH, MMNS).
Myriophyllum laxum Shuttleworth ex Chapman (Haloragaceae). Neither
Lowe (1921) nor Jones (1975) includes it for Mississippi, but PLANTS does.
Mississippi marks the species’ western range limit. This southeastern United
States endemic is often misidentified as M. pinnatum (Walter) BSP. or M.
heterophyllum Michaux.

Vouc

her specimens: Jackson Co.: submersed aquatic in pond, about 7 mi NNE of Ocean

SORRIE AND LEONARD, Mississippi vascular plant records oy
Springs, 18 Sep 1970, S.B. Jones, Jr. 20548 (GA). Lamar Co.: Mossy Pond, 3 mi W of
Purvis, 21 Jul 1973, E. Decker 62-6 (USM); about 3 mi W of Purvis, abundant in lake, 29
Aug 1973, K.E. Rogers 9224 (GA, NCU, NLU). Perry Co.: N end of unnamed pond NE of
FS. 321, Camp Shelby; eosuleton originally discovered by D. Wyrick, 11 Sep 1995, S.W.
Leonard 9060 (MMNS).

Parnassia grandifolia DC. (Saxifragaceae). Large-leaved grass-of-Parnassus
is known from the state only from the southeastern quarter, where first col-
lected in the late nineteenth century. Eakes (1989) cited two populations,
in Forrest and Stone Counties. The Camp Shelby inventory has verified 17
occurrences, plus an additional station on private property. a oa occur
in seepage communities at the base of slopes, in clayey soi

Voucher specimens: Pearl River Co.: at least 1( 0 plants in boggy drainage on N side of
Hillsdale Road, 3.7 mi W of I-59, or 0.5 mi SE of US 11, between Poplarville and Lumberton,
27 Apr 1996, 8. W. Leonard 9447 (MMNS). Perry Co.: several colonies from a few plants to
400 or more, in seepages that drain into Dickey Creek, Camp Shelby, SE of New Augusta,

2 Nov 1996, S.W. Leonard 9691 (MMNS),
Polygala leptostachys Shuttleworth (Polygalaceae). Lowe (1921) only lists
one record, from Harrison County. This species ranges from central Florida
to southwestern Georgia and southeastern Mississippi. Polygala leptostachys
is even more slender than its close relative, P. verticillata L., and the difficulty
of detecting it may explain the paucity of herbarium collecnions range-wide.

Voucher specimens: Forrest Co.: Camp Shelby, vicinity of Range 45, 0.5 mi NE of Lake
Janney, along hillside slope of road, 19 Jul 1996, J. MacDonald 9781 (IBE). Stone Co.:
se Miss. Forest Lands, diverse mesic uplands S of Wire Road and E of route 15, 20 Aug

997, B.A. Sorrie 9433 with R.G. Wieland (bas).
spares epihydrus Raf. (Potamogetonaceae). Jones (1974) omits this
species from Mississippi, but PLANTS includes it. Hellquist and Crow (1980)
specifically state “northern Mississippi” in their range description. In any
event, this pondweed 1s rarely collected in the state. The junior author found
it in dense beds at several road crossings of a gravelly, fast-flowing stream
in Lawrence County. In protected pools along the stream, the upper leaves
have weakly expanded blades to 5 mm wide, but none floating. This site
and those in two nearby Louisiana parishes (Thomas and Allen 1993), mark
the southernmost locations for the species.

Voucher specimen: Lawrence Co.: swiftly flowing water of Tilton Creek at SW end of
Tilton Road, about 0.4 mi from Sauls Valley Church; plants in clear water from a few cm
deep to 0.5 m, 28 Dec 1996, S.W. Leonard 9726 (IBE, MMNS)

Rhynchospora curtissii Britton (Cyperaceae). Kral (1996) maps only Jackson
County for Mississippi. This rare sedge is an East Gulf Coastal Plain en-
demic, known from fifteen counties range-wide.

Voucher specimens: Hancock Co.: 3.5 mi NE of Santa Rosa, just N of Dead Tiger Creek,
clearcut adjacent to hardwood hammock, clay soil, 23 May 1981, 8. McDaniel 25108 (FLAS).
This specimen was originally determined as R. pletantha (Kukenthal) Gale. Jackson Co.:

900 Stipa 18(3)
Ocean Springs, 1898, 8.ML. Tracy 4891 (GH, NCU), wet depression in sandy longleaf pine
savanna about 3 mi N of Ocean Springs, 20 Jul 1955, Channell 3577 (DUKE); scattered
populations in seepy powerline S of Semmes Road, about | mi W of Fort Bayou Creek, NE
of Ocean Springs, locally common, 15 Aug 1997, B.A. Sorrie 9410 (GH, MNNS, NCU,
VDB)

Rhynchospora decurrens Chapman (Cyperaceae). Reported herewith new
to Mississippi. This is a seldom collected species, although ranging from
southeastern North Carolina to south Florida and Louisiana. Gale (1944)
cites no Mississippi collections and PLANTS omits it. Rhynchospora decurrens
inhabits temporarily flooded depressions in swamps, often with Taxodium
and Nyssa.

Voucher specimen: Pearl River Co.: 2 mi W of Picayune, low woods, 9 Jun 1965, FH.

Sargent 8629 (DUKE, VDB).
Rhynchospora fascicularis (Michaux) Vahl var. distans (Chapman) Small
(Cyperaceae). Gale (1944) cites one Mississippi collection without specific
location. This is still a poorly known and seldom collected taxon, despite
distinctive achene morphology. It ranges from southeastern Virginia to south
Florida to southeastern Mississippi; disjunct to Bermuda.

Voucher specimen: Stone Co.: disturbed roadside seepage in clayey soil, Univ. Miss.

Forest Lands, uncommon, with Afetris aurea, Lachnocaulon anceps, \3 Jun 1997, B.A. Sorrie
9285 (GH, NCU).
Rhynchospora globularis (Chapman) Small var. pinetorum (Small) Gale
(Cyperaceae). Reported herewith new to Mississippi. Despite ample differ-
ences from other varieties of R. ¢/obularis, most authors have not recognized
var. pinetorum. The achene characters noted by Godfrey and Wooten (1979)
and Weakley (in prep.) are diagnostic. RAynchospora ae var. pinetorum
ranges from North Carolina to southern Florida and eastern Texas; also western
Cuba and Jamaica. This sedge often occurs on calcareous substrates.

Voucher specimen: Jackson Co.: on wet areas, Fountainbleau Point, P.O. Ocean Springs,
29 Apr 1954, D. Demaree 35016 (G
Rhynchospora harperi Small ips Reported herewith new to
Mississippi. Kral (1996), and LeBlond (1997) do not credit this species to
the state. Knowledge of the distribution of this species has increased dra-
matically during the past several years (LeBlond 1997). This sedge occurs
exclusively in seasonally ponded depressions on the Atlantic and Gulf Coastal
Plains.

Voucher specimens: Jackson Co.: Cottonmouth Savanna, N side of Frank Snell Road, E
of route 613, common in pond cypress-slash pine depression in flatwoods, 10 Nov 1997,

A. Sorrte 9632 (GH, MMNS, NCU); same place, 19 Jun 1998, B.A. Sorrie 9786 (FSU,
IBE, VDB
Rhynchospora nitens (Vahl) Gray (Cyperaceae). Lowe (1921) and PLANTS

es)

SORRIE AND LEONARD, Mississippi vascular plant records 901

do not list it for Mississippi, but there are specimens at VDB from Forrest
and Jackson Counties (Kral pers. comm.). RAynchospora nitens is widespread
on the southeastern United States coastal plain, probably overlooked in
Mississipp1.

Voucher specimen: Forrest Co.: Camp Shelby, NE of McLaurin, os wet area along
and above Hartfield Creek, 15 Oct 1994, C.T. Bryson 14511 et al. (ctb). George Co.: circa
8 mi W of Ala. state line on hwy. 98, 25 Sep 1979, K.L. Gordon 1495 u ith). Burris (IBE).
Harrison Co.: oo Island, small, seasonally inundated depressions, 7 Sep 1998, B.A. Sorrie
9953 IBE, NCU). Jackson Co.: Horn Island, 20 Jul 1894, S.M. Tracy 3925 (MISSA); NE
of US 90 and ve 63, Moss Point, open wet area and ditch along US 90 and frontage road,
1 Sep 1993, C.T. Bryson 12838 with Newton (ctb); dammed streamhead, now a Nyssa biflora-
Eleocharis equisetoides-Scirpus etuberculatus dominated pond, W side of FS 404, 0.25 mi N of
Larue see De Soto NF, 1 Oct 1997, B.A. Sorrie eee with RJ. LeBlond (GH, MMNS,

U, bas). Lamar Co.: Mossy Pond, 3 mi W of Purvis, 1 Sep 1998, B.A. Sorrie 9929 with
SW. ee. Perry cc Camp Shelby, Leaf River Wildlife Management Area, T1S
ROW Sect. 22, 15 Oct 1994, C.T. Bryson 14533 with J. MacDonald (ctb).

Rhynchospora scirpoides (Torrey) Gray (Cyperaceae). Reported herewith
new to Mississippi. Like R. nitens, this species inhabits periodically inun-
dated sites, but is much rarer in the southeastern United States. Lowe (1921)
lists Pstlocarya corymbiformis Bentham, a synonym of R. scirpoides, from Horn
Island, but the specimen at MISSA is actually R. nétens.

Voucher specimens: Forrest and Perry cos.: frequent on moist sandbars along Black
Creek, E of Brooklyn, 28 Sep 1997, B.A. Sorrie 9569 and M. Pyne (GH, IBE, VDB). Jack-
so ay oe tor Pond (dammed streamhead), just NE of jet. of Wire Road and FS 404-A,
Uni orest Lands, two robust plants on ee pond bottom, 24 rice 997, B.A.
nas 7 Ses NCU). Lamar noe 2.1miNo mberton on US 1 andy clay of
seepage area around pine woods pond, 4 Sep 1965, 5 a 25854 Sie ee Pond, 3
mi W of Purvis, 1 Sep 1998, B.A. Sorrie 9930 with 8. W. Leonard (NCL
Rhynchospora tracyi Britton (Cyperaceae). Lowe (1921) reports this sedge
on the authority of Small, but gives no localities. The following will serve
to document its presence in Mississippi.

Voucher specimens: Jackson Co.: cypress pond between Ocean Springs and Gautier, 28
Jul 1955, R.K. Godfrey 53700 with R.B. Channell (FSU, GH); hwy. 90 just W of Ala. line,
wet soil at edge ie marsh, 26 Sep 1970, K.E. Rogers 4612-A (SMU), 4612-B (NCU);

ue in shallow water, no date, K.E. Rogers 4627-A (NCU); edge of Cladinm marsh
ae S 90, circa i mi N of Orange Lake, 11 Sep 1980, K. Gordon 2255 (ctb, MMNS),
ommon in disturbed cypress depression at junction of Nut Bank Road and connector to
route 613, about 3.5 mi SSW of Big Point village, 2 Oct 1997, B.A. Sorrie 9588 with R.J.
LeBlond (GH, IBE, NCU).
Ruellia pedunculata Torrey ex Gray ssp. pinetorum (Fern.) R.W. Long
(Acanthaceae). This seldom collected taxon is vouchered from about 25 counties
range-wide (South Carolina to panhandle Florida to western Louisiana); half
of which are concentrated in a small area from Washington County, Ala-
bama to Tangipahoa Parish, Louisiana. Previously, this plant had been col-

902 Sipa 18(3)

lected in Harrison and Wayne Counties (MSNHP); here we add four more
counties. Because it is difficult to detect when not in flower, we suggest
that R. pedunculata ssp. pinetorum is probably not as rare in Mississippi as
previously assumed.

Voucher specimens: Forrest Co.: frequent . pine savannas in Ranges 43 and 45, Camp
Shelby, - May 1998, 8.W. Leonard 9954 (MMNS). Hancock Co.: Stennis Space Center,
NASA, 13 km SE of Picayune, pipeline at mi : off main canal, W of spillway, 29 Apr
E007: 7 Wooten s.n. (USM). Marion Co.: oe aang in powerline right-of-way through
low grounds, 1.2 mi N of hwy. 35 on E side of hw 3, 5 Sep 1998, 8. W. Leonard 10085
(MMNS). oy River Co.: about 5.5 mi N of Mc ai aoe 18 Aug 1980, 8. McDaniel
24337 (FS equent in A ae roadside 0.2 mi E of jet. of MS 26 and MS 43, 30 Apr
see SW. oe 9928 (MMNS). Perry Co.: about 30 ane in moist pine plantation on
side of Dickey Creek and E of North Tank Trail, Camp Shelby, 1 , 1998 - W. Leonard
ae (MMNS); burned-over hillside drains of Impact Area oe o N of South Tank
Trail and 1.0 mi E of Middle Creek, Camp Shelby, 11 May 1998, S. W. see (MMNS).

SEE

—

z

Sagittaria isoetiformis J.G. Smith (Alismataceae). Reported herewith new
to Mississippi. This is one of the more difficult species of Sagittaria to find,
because plants are often obscured by dense colonies of Eleocharis robbinsit,
Leersia hexandra, or other aquatics. Two locations are known from Camp
Shelby, a range extension westward from Covington County, AL.

Voucher specimen: Perry Co.: mostly subme - -d and not abundant in dense vegetation

fringing gum pond behind Mars Hill Church, Camp Shelby, 20 Sep 1994, J. MacDonald
7728b with D. Wyrick (IBE); same place, 8 Sep 1995, 5. W. aie 9057 with J. MacDonald
(MMNS), J. MacDonald 9107 (IBE).
Scleria reticularis Michaux (Cyperaceae). Reported herewith new to Mis-
sissippt. We here treat S. reticularis in the strict sense, without var. pubescens
Britton (= S. muehlenbergii Steudel). The two warrant treatment as full spe-
cies, based on morphological characters, habitat, and distribution, as treated
by Weakley (in prep.). Scleria reticularis sensu stricto ranges discontinuously
on the coastal plain from southern New Hampshire to the Florida panhandle
to Mississippi, disjunct to Michigan and Indiana. Itc is very rare south of
the Carolinas. Scleria muehlenbergii ranges continuously from southern New
Jersey to south Florida, west to Texas and Oklahoma, with scattered records
inland, in the West Indies, and in Central America. It is abundant in the
southern Atlantic and Gulf Coastal Plains.

Voucher specimen: Forrest Co.: hwy. 49 ca. mi N of Stone County line, shallow
pond W of hwy. across fence, 1970, K.E. Rogers aa: B (VDB), 4660-C (NCU). Lamar
Co.: Mossy Pond, 3 mi W of Purvis, uncommon on sandy-peaty shel fon NW shore, | Sep

1998, B.A. Sorrie 0936 with §.W. Leonard (GH, IBE, NCU).
Sorghastrum apalachicolense D.W. Hall (Poaceae). Shortly after this species
was named and described from Florida (Hall 1982), S. McDaniel found it
in Lamar Co., Mississippi (specimens at FSU, IBE) and K. Gordon found it

SORRIE AND LEONARD, Mississippi vascular plant records 903

in Pearl River County (MSNHP database). Until our survey work, Sorghastrum
apalachicolense was thought to be a rare disjunct in Mississippi, but it now
has been documented from nine counties. Moreover, this grass is quite common
in suitable habitat: frequently burned, mesic longleaf pine-oak-bluestem
communities with diverse herbaceous layers. We observed several thousand
plants in 1997 and 1998.

In 1997, SWL spent many hours observing the inflorescence develop-
ment of S. apalachicolense, S. secundum (E\.) Nash, and S. e//iottii (Mohr) Nash,
species often misidentified on specimen labels and annotations. Because a
the arching-drooping panicle axis, even soon after breaking the sheath, S.
elliottii differs markedly from the other two, in which the axis is straight to
slightly arching. Furthermore, S. e//iotti7 flowers a month later than S. apalachicolense

(in southern Mississippi) and about two weeks later than S. secundum (in
southern Alabama).

As the inflorescence emerges from the apical leaf sheath of 8. apalachicolense,
it takes the form of a compact plume with mostly ascendant branches. The
branches and branchlets diverge in an irregular pattern as anthers reach maturity;
later, as seeds develop and are shed, the branches return to the ascendant
position. In S. secundum, the branches and branchlets diverge in a very regu-
lar manner so as to form a secund inflorescence in which the florets line up
uniformly along one side of the axis. Each floret appears equidistant from
each other and from the axis. After seed maturation, the branches return to
the ascendant position. Thus, specimens taken prior to full anthesis, or to-
ward the end of seed dispersal, are superficially similar in these two species.
In addition to the spikelet width character given in Hall (1982), 8. apalachicolense
specimens possess a short collar of bristles at each panicle branch node; these
bristles are greatly reduced or absent in S. secundum.

Our observations of Sorghastrum reveal other important points. First, S.
secundum is absent from Mississippi; specimens so labeled from Lamar County
{S.B. Jones, Jr. 2460 (FSU)} are S. apalachicolense. In fact, we are unaware of
any verified record of S. secundum from west of the Mobile-Tensaw Delta,
Alabama. Second, flowering of 8. apalachicolense occurs about three weeks
later in Mississippi (late August to early September) than in the Florida
panhandle. We can offer no explanation for this discrepancy. Thus, in Florida,
flowering and fruiting of S. apalachicolense overlaps no other species, but in
Mississippi, it overlaps the onset of S. wwtans (L.) Nash. Third, the very short
life cycle—maximum of four weeks from breaking sheaths to seed set—
gives S. apalachicolense a short detection window.

Voucher specimens: Forrest Co.: longleaf pine woodland, 1.0 mi WSW of Grapevine
Road along FS 310-B, Camp Shelby, 6 Sep 1996, 8.W. Leonard 9622 (MMNS). George
Co.: mixed pine-hardwood stand N of Walker Road, Univ. Miss. Forest Lands, 22 Sep 1997,

904 Stipa 18(3)

B.A. Sorrie 9518 (US, bas). Harrison Co.: frequent in powerline right-of-way in sandy
soil, about 1 mi NE of Lizana along Saucier-Lizana Road, 7 Sep 1997, S.W. Leonard ae
(swl); rare in mesic longleat/slash pine woods by CC Camp Road Bog, S side of rou e 43
between Lyman and Lizana, 27 Sep 1996, B.A. Sorrie 9069 with R.J. LeBlond (bas). Jack-
son Co.: uncommon in multi-age longleaf ae -oak uplands N of Larue Road about 0.5
mi NW of Old Biloxi Road, De Soto NF, t 1997, B.A. Sorrie s.n. with R.J. LeBlond
(bas); cut-over pinelands 0.3 mi S of Wire se - on east side of hwy. 57, 7 Sep 1997, S.W.
Leonard 9879 (MMNS). Lamar Co.: very large population in thinned longleaf pine-bluestem
upland along Dobson Road, | Sep 1998, B.A. Sorrie 9938 with 8. W. Leonard ee IBE,

US); same place, 4 Sep 1998, B.A. Sorrie 9944 (BRIT, GA, GH). Marton Co.: infrequent
in pinelands along hwy. 13, 0.6 mi NW of Lamar Co. He 5 Se P 1998, S.W. ee 10082
(MMNS). Perty Co.: scatte nee in longleaf pine woods S of pipeline on both sides of FS

327, 3.2 mi S of FS 309, Camp Shelby, 8 Sep 1996, § a ete 9052 with J. MacDonald
(IBE). Stone Co.: frequent in mesic well-burned ce ees sine uplands, Univ. Miss. Forest
Lands, sheaths Rc and inflorescences opening 21-23 Aug 1997, spikelets nearly all
dropped by 23 Sep, B.A. Sorrie 9444 (MMNS, bas), 9446 (G H, NCU, bas), 94517 (IBE),
9452 (NCU, US), 9454 (DUKE, FSU, GH), 9533 (US).

Spiranthes brevilabris Lindley var. floridana (Wherry) Luer (Orchidaceae).
Reported herewith new to Mississippi. In a treatment of the Spiranthes of
the state, Morris (1989) could find no documentation of its occurrence, but
suggested that S. brevilabris and its var. floridana might occur. Although
the variety floridana supposedly ranges from North Carolina to Florida and
Texas, current information on distribution and abundance is virtually non-
existent. Most botanists have never seen this orchid and the Florida Natu-
ral Areas Inventory (the state with the most historical records) does not track
it. Knowledge of the range and current abundance of the nominate variety
is even poorer. Furthermore, confusion with S. /acera (Raf.) Raf. var. gracilis
(Bigelow) Luer has clouded specimen records, literature reports, and range
maps, including those of Luer (1972). Observations in 1997 suggest that
this orchid appears in the first growing season following a fire, when com-
petition from coarser herbs and shrubs is at a minimum. Plants mature and
senesce in only a few weeks, providing a brief window for detection. Mis-
sissippi populations all support less than twenty individuals. The Jackson
County specimen at AMES was originally determined as S. gracilis, and is
mounted on a sheet with other specimens of that taxon from Connecticut,
Illinois, and Vermont.

Voucher specimens: Harrison Co.: lobe of large seepage bog near FS 424, De Soto NF,
burned in winter/spring 1997, 22 May 1997, B.A. Sorrie 9251 (AVES). Jackaon Co.: Ocean
Springs, 29 April sa ES. Earle s.n. (AMES), seepage bog near FS 409-E, De Soto NEF,
burned winter 1997, 16 Apr 1997, B.A. Sorrie 9157 (bas). Stone Co.: mesic longleaf pine-
oak-grass upland a hardwoods mostly removed, very diverse herb layer, E of route 15
and S of Wire Road, as Miss. Forest Lands, 16 May 1997, B.A. Sorrie 9231 with R.G.
Wieland (MMNS, NCU
Stylisma aquatica es Raf. (Convolvulaceae). PLANTS lists this spe-
cies for Mississippi, but Myint (1966) does not map it there. This, our only

SORRIE AND LEONARD, Mississippi vascular plant records 905

pink flowered Sty/isma, inhabits intermittently ponded depressions, often
with Taxodium ascendens and Nyssa biflora in the canopy.

Voucher specimens: Harrison Co.: locally common ia seasonally ponded depression in
pine neon off of FS 452, S of Wortham Road, De Soto NF, grass-sedge dominated,

with Dich rectifolium, D. wrightianum, Lycopodiella alopecurotdes, 27 Sep 1997, B.A.
Sorrie 9558 with S$. W. i enadendl Moore (MMNS, NCU). jack con on disturbed cypress
depression at jct. of Nut Bank Road and connector to route 613, about 3.5 mi SSW of Big

Point village, abundant, 2 Oct 1997, B.A. Sorrie 9586 with RJ. LeBlind (GH. IBE, NCU);
Cottonmouth Savanna, N side of Frank Snell Road, E of route 613, cypress-bordered de-
pression in pine flatwoods, 1 Oct 1997, B.A. Sorrie with R.J. LeBlond 9582 (GA, DUKE).
Perry Co.: frequent in and around a wet weather pond on E side of FS 322, 0.5 mi N of FS
307 in Camp Shelby Training Area 63, 4 Jun 1996, S.W. Leonard 9529 (MMNS).

Stylisma pickeringii (Torrey ex M.A. Curtis) Gray (Convolvulaceae). Lowe
(1921) and Myint (1966) do not include this species for Mississippi, but
PLANTS does. The subspecific disposition of Mississippi specimens is prob-
lematic—characters they possess do not fit the published descriptions or
the key provided by Myint (1966). Supposedly, S. pickeringii var. pattersonii
(Fern. & Schub.) Myint can be separated from var. pickeringii by style branches
1.0—1.5 mm long vs. 2—3 mm, style a definitely unequal vs. mostly
equal or slightly unequal, stylopodia 1-2 mm long vs. up to 3—4 mm, and
sepals mostly acute or acutish vs. mostly obtuse. In Mississippi plants, style
branches are all less than 0.7 mm and vary from equal to definitely unequal,
stylopodia are all less than 2 mm, and sepals vary from acute to obtuse. The
resolution of this problem must await more detailed studies. The nominate
variety ranges discontinuously on the coastal plain from New Jersey to central
Alabama; var. pattersonii ranges from western Louisiana and southern Ar-
kansas to eastern Texas, Kansas, and Iowa. The following document the known

extant populations.

Voucher specimens: Forrest Co.: ce in extensive sand deposits in old floodplain
of Leaf River, N of Faulkner Street and E nes Street Baptist Church, Hattiesburg, 18

Jun 1998, B.A. Sorrie ied with §.W. ae GH, MMNS, NCU). Perry Co.: rare
in white sand in cut-over longleaf pine scrub behind Mars Hill Church, Camp S ple
Oct 1994, J. MacDonald 7953 with C.T. Bryson (IBE); same place, 10 Jul 1996, S. W. Leonard
9556 (MMNS); same site, 25 Sep 1997, B.A. Sorrie 9549 (bas, NCU). Wayne Co.: xeric,
deep white sands of hills along Brewer Town Road, De Soto NF, uncommon, with Pinus
palustris, Quercus laevis, Aristida tuberculosa, Rhynchospora megalocarpa, 26 Sep 1997, B.A. Sorrie
9551 with M. Pyne (bas).
Tridens carolinianus (Steudel) Henrard (Poaceae). Lowe (1921) does not
include this species (nor under any synonym), but PLANTS does. This striking
grass ranges from southeastern North Carolina to northwestern Florida to
western Louisiana, apparently nowhere common.

Voucher specimens: George oo: anssic oe as sane tore N of Walker Road, Univ

Miss. Forest Lands, rare, with Ti 2, S. nutans, Peal
bifidum, P. floridanum vat. a Aristida ies 22 Sep 1997, B.A. Sorrie s.n. (bas).

906 Sipa 18(3)

Pearl River Co.: Poplarville, 1932, H.R. Reed 324 (FSU), cepa originally determined
as Triodia drummondii Scribner & Kearney; McNeill, 23 Sep 1931, H.R. Reed 147 (FSU),
specimen originally det aie as Triodia elliottii Bush. Stone Co.: mesic longleaf-oak upland
N of Wire Road in Henley Park, Univ. Miss. ious Lands, uncommon, associated with high
a of SS and herbs, 23 Sep 1997, B.A. Sorrie 9535 5, NCU). Wayne
Co.: De by FS 245, ae in 1996 and
1997, ne ene herbaceous layer, 26 Sep 1997, B.A. Sorrie s.n. and M. Pyne (bas)

=

» NF, rare in mesic longleaf pine- oan nelend

Utricularia olivacea C. Wright ex Grisebach (Lentibulariaceae). New to
Mississippi. This neotropical species occurs in the United States as scat-
tered populations on the coastal plain from New Jersey to Florida. The plant's
extremely small size, even when flowering, makes detection difficult. Moreover,
U. olivacea is sensitive to changes in water level, so that it may not appear
annually at a given site. The Hinds County specimen names three lakes, at
least one of which was a natural arm of the Pearl River (R. Wieland, Mis-
sissippi Natural Heritage Program, pers. comm.). The 1935 date represents
the second oldest collection in the United States and the first outside of
Florida.

Voucher specimens: Hinds Co.: “Spring Lake; Clinton Lake; Hederman Lake”, 1935—
1945, EF Cook s.n. (MMNS). Jackson Co.: ponded gum swamp at FS 404, 0.25 mi N of
Larue Road, DeSoto NF, common in very shallow water and becoming Srances with U,
biflora, 18 Aug 1997, no sign of any sie on | Oct 1997, B.A. Sorrie 9416 (GH, NCU,
FSU, MISSA).

ACKNOWLEDGMENTS

We wish to thank the curators of the following herbaria for making available
their collections and facilities: AMES, DUKE, FLAS, FSU, GA, GH, IBE,
MISSA, MMNS, NCU, US, USM, VDB. Doug Goldman of the University
of Texas kindly provided several specimen records for Calopogon multiflorus.
Ken Gordon and Ron Wieland of the Mississippi Natural Heritage Pro-
gram made available database records. Charles Bryson of the US Depart-
ment of Agriculture in Stoneville, MS, Wayne Morris of North Georgia State
University, Dahlonega, and John MacDonald of Mississippi State Univer-
sity, Starkville, reviewed previous drafts and provided specimen data for
several taxa. Julie Moore provided logistical and technical support, as well
as coordinated the Camp Shelby and UMISS surveys. Alan Albritten pro-
vided lodging and technical support on the University of Mississippi For-
est Land. Publication costs were covered by the Mississippi Military De-
partment, National Guard Bureau, Department of Army. To all we extend
our sincere thanks.

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~
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DOCUMENTED CHROMOSOME NUMBERS 1999:1.
CHROMOSOME NUMBERS AND POLLEN
DIAMETER VARIATION
IN THELESPERMA (ASTERACEAE)

LEE F. GREER! and A. MICHAEL POWELL

Department of Biology
Sul Ross State University,
Alpine, TX 79832, U.S.A.

ABSTRACT

Diploid and tetraploid chromosome numbers and pollen measurements are reported for
95 collections from five taxa of Thelesperma (Asteraceae). These ct unts include
the first 17 diploid (2x) reports for T. megapotamicum var. ambiguum. A gor les 6k between
ploidy level (2x, 4x) and pollen diameter (mm) is found in diploid and polyploid plants of
T. megapotamicum and T. simplicifolinm. In both taxa, multivalent configurations (IV, V, VI,
X) are found. A recurring oversized bivalent (II) was eee in up to 23 of the 103 col-

lections from four of the five taxa studied.

RESUMEN

li -ontaron ntimeros cromosomaticos ea oides sy ietiae sides, »Y medidas

de polenice oe colecciones, Ths cuales en total agruy de Thelespe
Lo incluyen 17 de lelipoides me de T. mgaoranican vat. a oe
jae

publicados ahora por primera vez. En plantas poliploic eT. megapotamicum y
T. simplicifolinm se Paha una derelaaen entre el nivel de ploidia (2x, 4x) y el didmetro
del polen (mm). Ademds ambos téxones presentaron configuraciones multivalentes (IV, V,
VI, X). ants en 23 de las 103 colecciones se observ6 una recurrencia de bivalentes
(II) con tamafio incrementado, en cuatro de los cinco taxones.

Melchert (1963) included approximately 169 documented chromosome
counts for 10 taxa of Thelesperma. Prior to the Melchert (1963) study, chro-
mosome numbers were known for 10 of the 15 taxa recognized in his treat-

ent, and counts of 2 = 8,9, 10, 11, 12 and 22 were already established for
Thelesperma. The pre-1963 counts led erroneously to the notion that every
taxon of Thelesperma was characterized by a single chromosome number. Melchert
discovered 2x dysploid series in four taxa of Thelesperma and dysploid dip-
loid and polyploid plants in three taxa. The documented counts reported
by Melchert (1963) were never published. About 27 chromosome counts
for Thelesperma published in 15 papers since 1966 (see Goldblatt & Johnston

'Correspondence should be addressed to L.Greer, Loma Linda University, Dept. of Nat'l.
Sci.-Biology Section, Loma Linda, CA 92350, U.S.A., lgreer03¢@ns.llu.edu.

Sipa 18(3): 909-925. 1999

910 Sipa 18(3)

1990, and earlier Indices to Plant Chromosome Numbers) also revealed
chromosome number variation in the genus. Certain meiotic irregularities
in Thelesperma, mostly one to five rings-of-four in diploids and polyploids,
were reported by Melchert (1963), Strother (1976), Keil and Pinkava (1976),
and others.

We document numerous chromosome counts from five taxa of Thelesperma.
Most of the counts resulted from a populational study of the diploid-tetra-
ploid T. megapotanicum-T. ambiguum complex, which was the focal point of
thesis research (Greer 1997). We comment on meiotic irregularities and
the correlation between ploidy level and pollen diameter in Thelesperma.

METHODS AND MATERIALS

Capitula for meiotic analysis were fixed in Modified Carnoy’s Solution (4
chloroform: 3 ethanol: | glacial acetic acid; v:v:v). Standard squash tech-
niques (Turner & Johnston 1961) and acetocarmine stain were employed in
chromosomal observations. The sources of pollen for pollen size compari-
sons were the same as those for meiotic analyses. Pollen sizes were mea-
sured by ocular micrometer calibrated with slide micrometer such that 1
ocular unit equaled | Um (Greer 1997). Often up to 10—30 pollen grains
were measured for a given collection.

Correlations between pollen diameter and ploidy level were established
by a two-step process. A consensus measurement was estimated for the most
frequent ranges of pollen diameters found in each voucher collection (in-
clusive and exclusive of the pollen exine, 1.e., mm exinate and intinate).
These consensus measurements (in mm exinate ail intinate) were then plotted
against the inferred ploidy levels of the respective specimens (Fig. 1).

All the voucher specimens are preserved at the Sul Ross State University
Herbarium (SRSC) except for the B.L. Turner collections of T. filifolium vat:
filifolium, which are housed in the Herbaria of the University of Texas at
Austin (TEX and LL).

RESULTS

Included are 95 original chromosome number reports for four species (five
taxa) of Thelesperma. Specimen citations are accompanied by meiotic chro-
mosome numbers, configurations, irregularities, and pollen diameters in

Table 1. Certain capitulum characters of T. megapotamicum vat. megapotamicum
and 7’. megapotamicum var. ambiguum are included, e.g., heads radiate or dis-
coid and disc corolla coloration, because the putative hybrids of these taxa
are thought to be distinguishable by variation in these characters (Melchert
1963; Greer 1997). It is important to note that both 2x and 4x chromo-
some numbers and pollen grains with variable diameter were often observed

GREER AND POWELL, Chromosome numbers in Thelesperma ou

Pollen diameter - Ploidy level

60
n 50 ‘
“
o
E
e)
=
2
&
&
2 , Exinate
o
; = a Intinate
an)
ay — 5 per. Mov. Avg.
(Exinate)
9) |
1x 2x 3x?

Ploidy level

Fic. 1. Plot of exinate (small diamonds) and intinate (small squares) pollen diameter con-
sensus Measurements versus ee ative ploidy levels a 60 voucher collections, Table 1): 1
pollen measurements (mean = 21 mm, s.d. = 1.23); 2x pollen measurements (mean = 28,
s.d. = 2.7); 3x pollen measurements (m = 42, s.d. = a Trend line follows a five percent
moving average for exinate diameters.
in the same populations and even in the same individual plants (Table 1).
The Turner collection of T. fil/ifolinm var. filifolinm in Kimbel County yielded
the rare euploid cytotype of z = 11 (Greer 1997). Most collections of T.
filifolinm var. filifolinm have the dysploid cytotypes of 7 = 9 and = 8 (Melchert
1963). This chromosome number (Greer 1997) confirms the continued existence
of this rare cytotype in Kimbel County which was first reported by Melchert
(1963).

DISCUSSION

The 17 diploid (27 = 22) chromosome counts obtained for T. megapotamicum
var. ambiguum (RESULTS) represent the first diploid (2x) chromosome numbers
reported for the taxon. Previous reports (Melchert 1963; Turner & Flyr 1966)
suggested that var. ambiguum was exclusively tetraploid throughout its range.
Melchert (1963) recognized T. megapotamicum and T. ambiguum as distinct
species. He listed 42 counts for the complex, 17 diploid and tetraploid counts
for I’. megapotamicum, \9 tetraploid counts for T. ambiguum, and six tetraploid

912

TABLE |
|

. The meiotic chromosome pecan cont gunetions, irregularities, and pol
10

and loce at

Stipa 18(3)

s with
tire ey eee | . 1]

by en Observed al heal configurations and both
cluded. The putative presence of he oversized bivalent is designated
from more than one plant are indicated as (poy sulational). Authorities for the

pollen measurements

taxonomy are noted below!.

Oo f <
exinate and intinate pollen diameters are in-

oy 71), Chromosome counts or

Taxon & Locali Ploidy Chromosome — Pollen diameter (Lm)
capitular characters collection no. level number (exinate & intinate)
(w=11)
T. filtfolinm var. filifolium Tex.: Harper County = 2x 2n = OIL UD)
Red-brown disc, radiate 9 Tvrner s.n. (a,b) 2n = LOH UI?)
T. filifolinm var. filifolinm Tex.: Kimbel County = 2x n= 11 (Anaphase I: Only one
Red-brown disc, radiate = Turner s.n. side countable)
T. longipes Tex.: Crockett County =2x 18-20-24 Um exinate
Yellow discoid 5 mi N of Ozona 14-16-18 Um intinate
Greer 122a,b =4x 26-28 Lm exinate
(populational) 22-23 Lum intinate
T. megapotamicum Tex.: Terrell Coun 2x 2n = 111]
var. megapotamicum 5S mi E of Longfellow
Yellow disc, radiate Powell, 267 le
(red-brown 75%, orange (populational)
& yellow, ca. 12% each)
T. megapotamicum Tex.: Brewster 2x 20—23(—25) Um exinate
var. megapotamiicum County ~11.6 mi NE 16-18(—22—24) um
Yellow discoid of Alpine intinate
Greer 111 =4x, 6x? 29—30(—45) Lum exinate
26-29(-40) Um
intinate
T. megapotamicum 7 Brewster County 2x 2n = 111
var. megapotamicum ni S of
Yellow discoid, reddish — Fr. ae
disc lobes (red-brown at Powel! 2735
maturity)
T. megapotamicum Tex.: Pecos Count 2n= 110 GU!) 18-20 Um exinate
var, megapotamicum 3 mi E of TX 1776) 11 (10—-)15—18 Um
Greer 114 =4x, Ox intinate
30 [lm exinate few to
40 Um exina
T. megapotamicum NMex.: Socorro 2x 2n = 1111
County~34 mi E

var. Weve i une

Yellow disc

of San Antonio
owell .

2528

'Taxonomy with anenoee>: Thelesperma Lessing; 1. filifolium (Hook.) A. Gray var. filtfolinm;
T. longipes A.C tze var. megapotamicum, T. megeapotamicum
(Spreng.) O. Kuntze var. sian th, Gray) Shinners; 7. s#mp/icifolinm A. Gray var. simplicifolinm.

-
Cc

xray dejan I (Spreng )

GREER AND POWELL, Chromosome numbers in Thelesperma

13

Taxon & ocality & Ploidy Chromosome Pollen sos laa
capitular characters collection no. level number (exinate & intinate)
(x=11)
T. megapotamicum Tex.: Hudspeth fv 2n = 2211
var. ae mace ae County 12.5 mi E of
Yellow discoid ell Cit
Powell 2831
Tex.: Presidio County 4x 2n= 1811 + 21V

T. megapotamicum

var. megapotamicum
Yellow disccoid (~ 50
red and yellow discs)

T. megapotamicum
var, megapotamicum
Yellow discoid

T. megapotamicum
var. a ra
Yellow discoid

T. megapotamicum
var. megapotamicum
Rich, golden yellow

ee ;

T: megapotamicum
var. megapotamicum
Yellow discoid

T. megapotamicum
var. ee ae

Yellow dis

T. megapotamicum
var. me gapotamicum
Yellow discoid

T. megapotamicum
var. megapotamicum

Yello

ow disc, radiate

T. megapotamicum
var. megapotamicum
Yellow disc, radiate

xR

32 mi S of Marfa
Powell 2522

Tex.: Brewster County 4

iWo aye
Sloan and Powell 2
Tex.: Brewster aoe
W side of Alpin
Greer 96

: Brewster oie
W side of Alpin
Greer 100

Tex.: Brewster County
~10 mi NE of Alpine
Greer 110

Tex.: Pecos County
2 mi E of Imperial

reer 116a,b,¢
(populational
Tex.:
12.9 mi E of Imperial
Greer 119

wm

Pecos Count

Tex.: Terrell ee
E ay of Loz

Cany

ie ao

Tex.: Terrell County
rt

a: i7l

&

=
5

v
hp
xR

4

R

=6x?

2n= 18IIl + 21V

= 1811 + 21V (2H?)

On = 221]

2n = 2111?

2n = 221]

2n = 1311 + 41V? UI?)

2n = 221] 25—30(—3 1) Lm exinate

Anaphase | 18—23(—28) Um

dicentrics intinate
Immat.: 10 Lm exinate
6 Lum intinate

2n = 20-2110 ()

2n = 20-2111 25—35(—49 crushed)
(?) i

ratio.

2n = 2211 23-24 Lm exinate

19—21mm intinate
a a0 ) Lim exinate
3 Um intinate

1-3 IV

2n = 111] 19-22 Lum exinate {~2x}

2n = 221] 29 Lm exinate
¢

25-
Ay)

=

914

Table | continued

Sipa 18(3)

Taxon & Locality & Ploidy Chromosome — Pollen diameter (mm)
capitular characters collection no. level number (exinate & intinate)
(N= im
T. megapotamicum Tex.: Ward C panty 2x 2n = JII+11V
var. megapotamicnm 8 mi N of Jee (?)C1)
1776/1450
Greer 175
T. megapotamicum : Ward County, 2x 2n = LINC!)
var. megapotanicum : 0 2 mi N of 2n = 121]? (?)
Monahans 2n = YD +11V
rreer 177
T. ii aaah Tex.: Winkler 2x 2n = 11H Avg.: 20 Lm exinate
var. megapotamicun County, TX 14.7 mi
oung appearing plants = N of Monahans 4x Avg.: 29-30 Um
ear perennials? Greer 179 exinate
T. megapotamicum Tex.: Winkler County 2x 2n= 111
var. megapotamicum 4.5 mi NE of Kermit
Greer 181
T. megapotamicum Tex. : Winkler County 2x 2n = LIT US) I“ head: Most 18-20
var. megapotamicum 14 mi NE of Kermit [lm exinate
sreery 182 (132-)14-15 bm
(populational) intinate
2n = O11 + 11V few clusters: 27—30
tlm exine
2n = SI] + 31V 18—20 Um intinate
{=4x?}
or OIL + 31V (2) 2" head clusters:
2n = 41 4 —32 Um
+ IVI exinate {=4x}
2n = 1111 or 91
1IV 3 shead: —35 um
exinate
25-30 btm
intinate
T! megapotamicnm Tex.: Winkler County 2x 2) I]
var. as a ae mi NE of Kermit 2n = 11ILUI)
ome? Gr 83 2n = OIL + LIV
(I year ie ) Gao Ax 2n = 1811 + 21V
T. megapotamicum Tex.: Andrews County 2x 2n = LIT US)
var. megapotanticum of Winkler-Andrews
Reddish involucres County line
Greer 185
T. megapotamicum Tex.: Andrews County 2x a = 1111
Var, megapotanlicum 24.3 mi NE of Kermit = OIL + 1IV
Greer 180a on = LOU? U1)
7. megapotamicum Tex.: Andrews County 2x 11

var. megapotamicum

24.3 mi NE of K
Greer 1866

Sermit

2n =
2n = LOI? UI)

GREER AND PoWELL, Chromosome numbers in Thelesperma oO

Taxon & Locality & Ploidy Chromosome Pollen di (mm)
capitular characters collection no. leve number (exinate & intinate)
(x=11)
T. megapotamicum Tex.: Brewster County 4x 2n = 1111 + 31IV+1X
var. megapotamicum 36.4 mi S of I-10, 2n = 18]1 + 21V
Yellow discoid US 67 2n = 1711 + 11V + 1VI + LVI (71?)
31 May 1995 2n= 1711 + 11V
Greer 187
T. megapotamicum Tex.: Brewster County 4x 2n = 2211 (?)
var. megapotamicum 9.6 mi + i Marathon
ellow discoid Greer
T. megapotamicum Tex.: Pecos County 2x (19-)21—22 Lm exinate
var. megapotamicum 40.5 mi E of Marathon |7~18 Um intinate
Yellow disc, radiate Greer 206
T. megapotamicum Tex.: Terrell County = 2x 2n = 1111
var, megapotamicum 41.7 mi E of Marathon
Orange-yellow disc, Greer 216

radiate; large trilobate
ray florets

T. megapotamicum oe Terrell Count 2x 2n = 111]

var. megapotamicum 2.4 mi E of M ae

Yellow disc, eae 217

reduced radiate

T. megapotamicum Tex.: Terrell County Ax? 2n = 2211 (?)

var. megapotamicum >46 mi E of Marathon

Yellow discoid Greer 222

T. megapotamicum Tex.: Terrell County — 2x 2n = III]
var. megapotamicum Sl mi : 7 Marathon

Dark yellow disc, Greer 22

minute ray florets

T. megapotamicum Tex.: Terrell County 1x 2n = 22K] (?)

var. sais aaa 52.4 mi E of Marathon
Yellow, red-cinted Greer 228
discoi id

T. megapotamicum Tex.: Terrell County = 2x 2n = 11II
var. megapotamicum 51.5 mi E of Marathon
Yellow disc, radiate Greer 234a
T. megapotamicum Tex.: Terrell County 2n =1111 (HI)
var. megapotamicum 51.5 mi E of Marathon
Yellow disc, radiate Greer 234b ix 2n = 221
2n = 2011 + 11V

T. megapotamicum Tex.: Terrell County In a2 25, 28-30, ~40 Um
var, meg ipotamitum 29.1 mi W of Lozier Anas I exinate
Yellow discoid Canyon >? {=4x, 6x?}

Greer 202 (populational) per melocyte

916

Table 1 continued

Sipa 18(3)

‘Taxon &
capitular characters

ocality
collection no.

I] di ( \

(exinate & intinate)

T. megapotamicum

var. me ee “um
Golden

very une radiate

T. megapotamicnm
é

at
reduced), yellow style
branches
T. megapotamicum
var. me, Libya “um
Yellov c, radiate

(dark miata disc florets)

T. megapotamicum
var. megapotamicum
Yellow discoic

T. megapotamicum
var. megapotamicum
Dark yellow discoid

T. megapotamicum
var. megapotamicum

and var. ambiguum

Yellow discoid;

2 orange discoid

(one clump)

T. megapotamicum

var. ambiguum

Orange disc, radiate

T. megapotame mM

population of radiate
& discoid)

T. megapotamicum

var. ambiguum

Red discoid

Tex.: Terrell County
32.6 mi W of Lozier
Canyon

Greer 263

Tex.: Terrell County
32.6 mi W of Lozier
Canyon

Greer 204

Tex.: Terrell County

4.0 mi W of Crozier

ee ae

Tex.: Terrell County

~40+ mi W of Lozier

Canyon
Greer 27 1b
Tex.: Val Verde
County 5 mi W
of Del Rio

reer 155

Terrell County

A el

haa: populacional)

2n =

Tex.: Brewster
County 5 mi W of
Maratho

Powell oo

Tex.: Brewste

Saintes near es 1

Powell 2

x.: Brewster County

~17.1 mi NE of Alpine

Greer 113

=

SN
R

cad

=

2.4 mi E of Marathon

22-26, 40 Um
exinate
20-25 Um intinate

25—26(—30) Lim exinate

(16—)18—20(—22) Um
intinate

(up co 35 [tm intinate)

+ neric
Anaphase I: Separated
n= 1111+ 4IV

variable

GREER AND POWELL, Chromosome numbers in Thelesperma O17
axon & ocality & Ploidy Chromosome Pollendi (mm)
capitular characters collection no. level (exinate & intinate)
(x=11)
T. megapotamicum Tex.: Pecos County 2 2n = 111d)
var. ambignum 40.5 mi E of Marathon
Red disc, radiate; Greer 207
double layer ray florets
T. megapotamicum Tex.: Terrell County = 2x = 111
var. ambignum 41.7 mi E of Marathon
Red discoid Greer 213
T. megapotamicum Tex.: Terrell County = 4x 2n = 2211 (?)
var. ambignum 32.6 mi W of Lozier
range disc, radiate Canyon
(slender ray florets) Greer 265
T. megapotamicum Tex.: Brewster County 2x = 1811 + 21V
var. ambiguium Marathon
Red-brown discoid Powell and Powell 2540
T. gsm Tex.: Brewster County 4x 2n= 1811 + 21V
var. ambigu Marathon
Red- ein co Poul and Powell 2541
T. megapotamicum Tex.: Terrell County 26 2n = 11 UTI?)
var. ambiguum 51.5 mi E of Marathon
Orange disc, radiate; Greer 235 Zn 2210
some double ray florets (2 meiocytes)
T. megapotamicum Tex.: el County 4x 2n = 2211
var. ambiguium 10.2 W of Lozier 27 = 181 + 21V (I)
range disc, eee
ane Greer 255
T. megapotamicum Tex.: Terrell County = 4x 2n= 13— 19OIl + LIV
var. ambiguum 29.1 mi W of Lozier 2n = 22I1 (?)
Orange discoid almost: | Canyo
2 minute ray florets Greer 259
LT: megapotamicum Tex.: Terrell County = 2x 2n = 11K
var. ambiguum 0.9 mi W of Sanderson
Small red disc, radiate Greer 280
T. megapotamicum Tex.: Terrell County = 2x 2n = 11 (2-411)?
var. ambiguum 1.1 mi W of Sanderson
ed disc, radiate Greer 283
T. megapotamicum Tex.: Brewster County 4x 2n~ 1811 + 21V
var. ambiguum 70 mi S of Alpine
Reddish-brown disc, Powell and Powell 2540
radiate
T. megapotamicum Tex.: Terrell County 2x = 111

var. ambiguum
Red-brown disc, radiate

(red-brown 75%, orange

& yellow, ca. 12% each)

5 mi E of Longfellow
Powell, 267 1a

(populational)

918

‘Table 1 continued

Sipa 18(3)

‘Taxon & Locality & Ploidy Chromosome — Pollen diameter (mm)
capitular characters collection no. level number (exinate & intinate)
(w=I11)
T. megapotamicnm Tex.: Terrell County 2x 2n = 1111
var, ambigunm 5.5 mi E of Longfellow
Orange disc, ra al € ea) 26716
(red-brown 75%, orange (populational)
& yellow, ca. 120% each)
T. megapotamicum Tex.: Brewster County 4x 2n = 131l + 41V
var. ambigunm 36.4 mi S of I-10, Pn = 1511 + 1IV + 1X
Red discoid US 67
31 May 1995
Greer ies
T. megapotamicum ex.: Brewster County 4x 2n = 201 + 1IV
var. ambiguum 36.4 mi S of 1-10,
Reddish-tinged discoid = US 67
31 May L995
Greer 188)
di ca mmicune Tex.: Brewster County 4x 2n = 221]
var. ambiguum 12.9 mi f 2n = 2011
Red discot i US —
yellow style branches Greer lagging chromos.
(poy eee
T. megapotamicnm Tex.: Brewster County 4x 2n = 1811 + 21V
var. ambiguium 3.5 mi E of Marathon
Red discoid Greer 192
7. ies oasis Tex.: Terrell County 2x 2n= 1111
var. ambiguir 42.4 mi E of Marathon
Jeep orange disc, radiate Greer 218
T. megapotamicum Tex.: Terrell County — 2x 2n = IIIT (?) 20-21 Lm exinate
var, ambiguium >46 mi 7 of Marathon 15-18 [tim incinate
range discoid Greer 2n = 22II (?) 26 Lm exinat
(L pollen)
2n = 2211 20 Um intinate
(1 pol
SAN 30 Lim ate
(1 shattered pollen)
T. megapotamicum Tex.: Terrell County 2n= 2111 20-22, 24-26(-31)
var, ambiguum 52.4 mi E of Marathon 2n = 2211? Lim exinate
Reddish discoid Greer 229 (16—)20—2
intinate
2x, 4x}
7. eccrine Tex.: Terrell Cour 2 2n = 1111
var. ambiguum 52.4 mi - of Maré a
Orange Greer 2

large ray ig ae

GREER AND PowELL, Chromosome numbers in Thelesperma

Table | continued

19.

capitular characters

Locality &
collection no.

Ploidy
level
(w=11

Chromosome _ Poll li (mm

number (exinate & intinate)

T. megapotamicum

var. gicnne

Red discoid; some heads
more reddish than
others on the same plane

—

: megapotamicum
var. ambiguum

ed disc, radiate

T. megapotamicum

var. ambiguum
Orange disc, radiate
(large ray florets
some double layered)

T. me ein um

eae discoid,
orange een.
reduced ciate

T. megapotamicnm
var. ambiguum
Reddish discoid
LT. megapotamicum
var. ee
Rad

T. megapotamicum
var. ambiguum
Discoic

18 ee

var. ambignum

Discoid, orange style
ches

Tex.:
33.7 mi as of Marathon
Greer 2

a

Pecos County

—

Tex.: Pecos County
38.8 mi E of Marathon
201

: Pecos Co

Greer

- : mi oe of ae ee

Greer 2

Tex.: Terrell County
E side of Lozier
Canyon

reer 170

(populational)

Tex.: Maverick County
4 mi NW of Eagle
Pass Greer 144a.h.c.d

(popuational

—_

Tex. : Maverick County
W of

Tex.: Maverick oi
Rick ie ad
US 27

Greer ie

eX.! Mi werick County
WW of

Quemado
Greer 149

4

bx

a

2n = 221] (?)
2n= 1111
2n = 1I1II
2n = 221] 1°’ Head: 26—31mm
exinate
23-29 Um intinate
37-48 lm exinate
34, 42 Um intinate
2"d Head: ve
exinate
15-2] ss intinate
3" Head: 23 um
inate
19 fe intinate
28-31 [Lm exinate
23-27 Lm intinate
2n = 2 (?)

—_

two Anaphas
dicentric

2n = 2011

25—31mm exinate

(18-19-)23-30 [im

—36 Um (few)—39 Um

19-22 Lim exinate

11—18 Um intinate

20-31 [lm exinate

20-22, 25, 28 Um
intinate

920

Table | continued

Sipa 184)

& Locality & Ploidy Chromosome — Pollen diameter (mm)
capitular characters collection no. level ber exinate & intinate)
(v=11
=x? one cluster: 25-30
lm exinate
22-26 Um
intinate
T. megapotamicum Tex.: Val Verde adn 2n = 2211? UI) 28-29(—35) Lm exinate
var. oe County Del Rio, {6x?} Ac least | IV 19-20 [lm intinate
Red discoid Ss + II
Greer 151
T. megapotamicum ex.: Val Verde =2Nx 2n = 10-121 (?) 20-21 Um exinate
var, ambiguum County 5 mi W ae I 11—)15—18 Um
Orange disc, radiate of Del Rio dicentric intinate
Greer 15 lagging Immat.: 13-15 Um
chromos exinate
=AN 2n = 22
(one cluster) = 20—25(—32) Um exinate
20-22, 26-29 Um
intina
Immat.: 20-21 Um
w/o exinate
T. megapotamicum Tex.: Val Verde =2x? 2n = LIL () 21-20 Um exinate
var. ambiguum County 5 mi W (15-)18—20 Ltm
Red discoid of Del Rio intinate
Greer 153 in 2n = 221] 25-30 Um exinate
2n = 2011 20-26 [tm intinate
Ox? chia —40 [im exinate
8x? dicentr —<50 [im exinate
T. megapotamicum Tex.: Val Verde =4x 25—31(—35) bum exinate
var. ambiguum County 5 mi W —24(—29) um
Red disc, reduced of Del Rio intinate
radiate Greer 154
T. megapotamicum Tex.: Val Verde . ounty 4x 2n = 22II (?)
var. ambiguum 5S mi W of Del I
Red disc, radiate Greer 157a,b,c,
(populational)
T. megapotamicnm ‘Tex.: Val Verde County =4x 2n = 2211
var. ambiguum 2.0 mi W of Comstock
Red-orange disc, radiate Greer 162
T. megapotamicum Tex.: Pecos County 2 2n = LIKI
var. ambiguum 40.5 mi E of Mar athon 2n = 1OI (2)
Orange discoid Greer 204
T. megapotamicum Tex.: Brewster County 44 = I8Il + 2 1Vs

var. ambiguum
Reddish-brown disc
lobes, corolla throats

ellow, radiate

Terlingua Creek

Powell 250°

GREER AND POWELL, Chromosome numbers in Thelesperma

Table | continued

axon & Locality & Ploidy Chromosome Pollen Sian
capitular characters collection no. evel number (exinate & intinate)
(w=11)
T. megapotamicum Tex.: Brewster County 2x 2n = 111
var. ambiguum 5 W of Marathon
Red-brown disc, radiate Powell 2684
T. simplicifolinm Tex.: Val Verde 2x LOM or LL (dy)
var. simplicifolinm County25.7 mi W e II may two IIs
of Pecos River elise)
Greer 166
T. simplicifolinm Mexico: Coahuila, Ax 2n = 201
var. semplicifolium Powell, D. Patterson,
Yellow disc, radiate D, Ittner 1584
T. simplicifoltum Crockett County 2x 2n= 10M U1) (22-)24—25 Lim exinate
var. simplicifolinm mi E of Jet. TX 14-17 [tm intinate
ae 90 =4x? 2 exinate
Greer 127 20—21(—23) Lm
intinate
=Ox? 42? Lim exinate
37? Um intinate
T. simplictfolinm Tex.: Val Verde =2x Avg.: 20 exinate
var. simplicifoltum County23.8 mi W (10-12)15
of Del Rio intinate
Greer 158
T. simplicifolinm Tex.: Kinney County 2x 2n = 10H; 20-22 [lm exinate
var. simplictfolinm 21.1 miNWo 2n = 10? 13? 15-18 Lm intinate
Quemado =4x a few cells: 25,
Greer 150 31mm exinate,
~23 Um
T. simplicifolinm te Menard ae 2x = 1011
var. simplicifolinm mi : - TX 29/8 Vee I ie
yw disc, radiate ae chromosomes
Curvicarpum achene form one bivalent (J
T. stmplicifolinm ex.: Menard County =2x 18—20 Um exinate
var. simplicifolinm | mi E of TX 29/83
Yellow disc, radiate Greer 138 =6x? 35—36(A0) Um exinate
Curvicarpum achene form base of echinations:
30 Lm
T. simplicifoltum Tex.: Maverick County 2x 2n=11N GUI) 20-23(—29, 30) Um
var. simplicifolinm Rio ae alley, exinate
US 27 {=4x?} —17 [lm intinate
ao 147
T. simplicifolium Tex.: Terrell County = 2x 2n= 1111
var. simplicifolinm 0.2 mi W of Lozier 4x 2n = 201] 20-21-24 Lm exinate
Yellow disc, large Canyon (13-)16—20 Lm
overlapping ray florets Greer 243 intinate

922 Sipa 18(3)

Table | continued

axon & Locality & Ploidy Chromosome — Pollen diameter (mm)
capitular characters collection no. level number (exinate & intinate)
(v= 1
T. simplicifolium Tex.: Val Verde 2x 2n = 20(?) 21-22 Um exinate
var. simplicifolinm County 0.7 mi W ot 2n = LOI (G31?) 1 16 5 Lm intinate
Pecos River =4x 26—28(—29-30) Lm
Greer 103 exinate
22—24(—26) Um
inctinate

counts for putative hybrids between the taxa. Because of their remarkable
morphological similarity, identical meiotic cytologies (27 = 22, 44 with
multivalents), and the ease with which they apparently interbreed where
sympatric, I’. megapotamicum and T. ambiguum are best treated as varieties
(Greer 1997). The only reliable distinguishing characteristics are range, the
slightly reduced habit of var. ambiguum, and disc coloration. Disc florets of
var. megapotamicum are yellow and heads generally are discoid; disc florets of
var. ambiguum are red-brown and heads are generally yellow-radiate.

The reports (Table 1) for the T. megapotamicum complex provide an ex-
panded understanding of how diploid and polyploid distribution in the species
is structured, especially in Texas. Thelesperma megapotamicum var. megapotamicum
is almost exclusively diploid (27 = 22) with yellow, discoid heads in most
of its range, which extends from Nebraska, Colorado, and Wyoming west
to Arizona, south across Trans-Pecos Texas into central Mexico, and east to
northwest Texas (Melchert 1963). In the predominant portion of its range
which extends from south Texas and near the Rio Grande Valley west into
Val Verde County, T. megapotamicum var. ambiguum is largely tetraploid (20
= 44) with red-brown disc corollas and large, yellow ray florets. However,
west of the Pecos River, mixed diploid and tetraploid populations of var.
ambieuum occur from western Val Verde County through Terrell and Brewster
counties (Greer 1997). The floral morphology of these populations of var.
ambiguum vary from radiate with red-brown discs, discoid orange, radiate
orange, to discoid red-brown. Along this same axis, var. megapotamicum plants
are also often diploid (27 = 22) with yellow discs and ray corollas. Popula-
tions of var. megapotamicum elsewhere in the Trans-Pecos are mostly tetrap-
loid (2 = 44) with yellow discoid heads, although there are occasional
with yellow discs and yellow ray Boece often reduced in length.

Melchert (1963) interpreted plants from this area resembling both var.
megapotamicum and var. ambiguum as hybrids or intergrades between the taxa.
He found only tetraploids in this area. This region of apparently extensive
morpho

slants

—

—

ogical intergradation seems to mark the sympatric overlap of the

GREER AND POWELL, Chromosome numbers in Thelesperma PAs:

ranges of var. megapotamicum and var. ambiguum from western Val Verde County
west into Brewster County. The intergradation is so smooth and continu-
ous that morphotypes were difficult to assign taxonomically.

Origins of polyploidy. The origins of polyploidy in T. megapotamicum remain
uncertain. Variation in floral character combinations in the sympatric re-
gion, high fertility, and the absence of univalents or trivalents in meiotic
observations, suggest that alloploidy is more likely than autoploidy in var.
ambiguum (Greer 1997). Because of the lack of morphological differentia-
tion between diploids and tetraploids, however, polyploidy in the typical
yellow discoid var. megapotamicum seems more likely to be autoploid. For
similar reasons, autoploidy also seems more likely in other polyploid taxa
of Thelesperma such as T. simplicifolinm and T. longipes (Greer 1997).

Multivalent configurations have been observed at meiosis in diploid (Keil
& Pinkava 1976; Greer 1997; Results) and in tetraploid (Melchert 1963;
Strother 1976; Powell & Powell 1977; Greer 1997; Results) cytotypes of
both T. megapotamicum vas. megapotamicum and T. megapotamicum var. ambiguum.
Melchert (1963) reported that in tetraploids, two rings-of-four (IV) were
most common, and that one and three rings-of-four also were observed. In
this study, we report that widely variable numbers of apparent ring and
chain quadrivalents (IV), hexavalents (VI), and even decavalents (X) were
found in populations of var. megapotamicum and var. ambiguum (Greer 1997).
Anaphase segregation was essentially balanced and fertility remained high
in spite of the multivalents (Melchert 1963) and the occasional occurrence
of dicentric chromosomes (Table 1; Greer 1997).

Large bivalent. A persistent large bivalent (II) was observed in meiotic
preparations in up to 23 of the 103 collections (Table 1; Greer 1997), most
noticeably in preparations from diploid plants. Such an oversized bivalent
has been observed in T. megapotamicum var. megapotamicum, T. megapotamicum
var. ambiguum, T. simplicifolinm var. simplicifolium, and T. filifolium var. filifolium
(Greer 1997). An oversized bivalent is also plainly visible in Melchert’s (1963)
meiotic camera lucida figures of the above taxa as well as in T. /ongipes, T
filifolinm var. intermedium, and possibly also in T. burrideeanum (Greer 1997).
Melchert (1963) did not call attention to the large bivalent. In one collec-
tion of 1. s*mplicifolinm var. simplicifolium (Greer 137; Table 1), the chromo-
somes of an oversized bivalent lagged in anaphase I. The lagging and di-
centric chromosomes observed (Table 1) may be associated with the large
bivalenc. A large bivalent would be more frequently expected to lag and to
undergo paracentric inversions because of its length. The large bivalent may
be a result of a massively unequal reciprocal translocation that occurred early
in the evolution of Thelesperma (Greer 1997),

Pollen diameter variation. Pollen diameter size variations seem to reflect

924 Stipa 18(3)

the ploidy levels of the plants that produced them (Greer 1997). From dip-
loid plants (2x) of T. megapotamicum, the broad range of pollen diameters for
presumed |x pollen (1x gametophytes) was 17—26 lm exinate and 11—21
Um intinate. From tetraploid plants, the range of pollen diameters for pre-
sumed 2x pollen (2x gametophytes) was approximately 20-35 Um exinate
and 18—29 Um intinate. Rare pollen found in preparations with even larger
diameters, 30—50 Lim exinate and 25—42 Um intinate, have been attributed
tentatively to 3x pollen (3x gametophytes). (See Fig. 1).

Diploid plants of T. Chai occasionally were found to produce
unreduced 2x pollen (ca. 24—30 Lm exinate) along with the more abundant
1x pollen (18—24 Lm exinate). In Andrews and Winilet counties where var.
megapotamicum is known to occur only as a diploid, discrete clusters of larger
pollen were seen in meiotic preparations, suggesting that localized tetrap-
loid (4x) microsporangial tissues are producing clusters of unreduced 2x
gametophytes. Plants of both var. megapotamicum and var. ambiguum from
Brewster, Terrell, and Val Verde counties, where populations are typically
tetraploid, consistently produce pollen of both sizes (ca. 24—30 [tm; ca. 18—
22 Um) ina nearly 1:1 ratio of putative 1x and 2x pollen. In tetraploids of
T. megapotamicum larger than normal pollen (30-35 Um) were occasionally
observed, suggesting that tetraploids also may be producing unreduced
gametophytes (3x?, Fig. 1).

The same kind of ploidy level—pollen diameter variation was also ob-
served in T. smplictfolinm var. simplicifolinm, which has both diploid and tetraploid
cytotypes. The correlation between known chromosome numbers and pol-
len sizes suggests that in certain species of Thelesperma a given ploidy level
may predominate in a specific population or plant, while gametophytes of
different ploidy levels are being also produced (Greer 1997). Similar pro-
duction of unreduced gametes has been reported by Beaman (1957) in Townsendia
(Asteraceae) and Powell and Sikes (1975) in Perity/e (Asteraceae).

ACKNOWLEDGMENTS

We thank Sharon Yarborough, Assistant Curator of the Sul Ross State
University Herbarium (SRSC), for her help in dealing with specimens of
Thelesperma. We are grateful to B.L. Turner for contributing capitula of Thelesperma

and for discussing the essence of his systematic observations concerning 7 helesperma
megapotamicum and other species of the genus. We appreciate loans of speci-
mens from the University of Texas (TEX, LL) and El Instituto de Botanica
Darwinion (SI), San Isidro, Argentina. We also thank Cristian Carvajal and
Raul Esperante for their translation of the abstract into Spanish.

GREER AND POWELL, Chromosome numbers in Thelesperma 92)
REFERENCES
Beaman, J.H. 1957. The systematics and evolution of Townsendia (Compositae). Contr. Gray
erb. 183:1-151.

Go LpsLaTT, P. and D.E. JOHNSON (eds). 1990. Index of plant chromosome numbers (series
of monographs from 1975-1987). Missouri Botanical Garden, St. Louis, Missouri.

GREER, 1997. Origins of polyploidy in Thelesperma (Asteraceae). M.S. thesis, Su
State Universi, Alpine, Texas.

Ket, D.J. and D.J. PINKava. 1976. Cl axonomic notes for Compositae
from the ae States and Mexico. Amer. J. Bot. 6: ee

MELcHert, T.E. 1963. Systematics of the genus Thelesperma (A crasonomi and chemo-

Ross

taxonomic aa Ph.D. dissertation, University of Texas, Aus

Powel, A.M. and S.A. PoweLt. 1977. Chromosome numbers of gypsophilic plant s
of che Chin bon Desert. Sida 7:80—90.

PoweLL, A.M. and S.W. Sikes. 1975. On the origin of polyploidy in Perityle rupestris (Asteraceae)
Sci. Biol. J. 1:132-137. (Oct-Nov

STROTHER, J.L. 1976. Chromosome studies in Compositae. Amer. J. Bot. 63:247—250.

TURNER, B. and D. Fryr. ee Chromosome numbers in the Compositae. X. North American
species. er. J.B 3:24-33,

TURNER, 8B. ia and M. C. a 1961. Chromosome numbers in the Compositae. III.
Certain Mexican species. Brittonia 13:64—69.

yECIES

fans

926 Sipa 18(3)
BOOK REVIEW

Brian E. 8. GUNNING AND Martin W. Steer. 1996. Plant Cell Biology:
Structure and Function. (ISBN Somme: pbk.; 0-867 20-509-

| (hbk). Jones and Bartlett Publishers, 40 Tall Pine Drive, Sudbury,
MA 01776. 800-443-5000. $37.50. ie numbers, no page num-
bers)

—

This book replaces the classic work, “Plant Cell Biology-An Ultrastructural Approach,”

published in 1975. It includes concn made using the following microscopic techinques:

conventional light, phase contrast, heen interference contrast, fluorescence (includ-
ing fluorescence antibody and immuno-fluorescence), confocal laser scanning, transmis-
sion electron (TEM) and scanning electron (SEM). Many techniques — anticlonal
antibodies, serial optical sections, shadow casting, freeze-fracturing, freeze-etching, ultra-
thin, freeze substitution, and a variecy of ancillary ones, are explained in detail i in oe
as well as in the legends.

The book is comprised of 60 plates in an 8.5 x 12" format, on high-gloss paper, with
sometimes oversized labels for the sub-plates, and an accompanying text for each. Several
black and white drawings scattered throughout the text are original artwork, but are not
numbered. There is also one figure of a “generalised plant cell” that places the organelles
and ultrastructural features of our beloved tetradodecahedron in perspective. The 60 plates
are arranged in groups according to subject, and include the following sections: an intro-
ductory survey, nucleus, ona reticulum, ae apparatus and coated vesicles, vacuoles,
mitochondria, nucleic acids in mitochondria and plastids, plastids, microbodies, cytoskel-

eton, cell division, transport ee cells, vase ae tissue, the plant surface, plant repro-
duction, the plant as a multicellular organism and the index.

Like its predecessor, this book is an instantaneous classic and a MUST for anyone teach-
ing gene! ral biology or plant biology. The clear, concise text and fantastic images would
serve the needs of everyone from high school biology advanced placement students to ad-
vanced undergraduate biology majors taking a cell com course. | unhesitatingly recom-
mend this excellent bargain to everyone!—John_J. Pipoly 1

SALVINIA MOLESTA (SALVINIACEAE),
NEW TO TEXAS AND LOUISIANA

COLETTE C. JACONO

United States Geological Survey
LI20-NW Fist Si.
Gainesville, FL 32053, U.S.A.

Initially discovered at a schoolyard demonstration pond in southeastern
Texas, the Federal noxious weed Sa/vinia molesta Mitchell has recently been
found in abundance at Toledo Bend Reservoir. This 75,300 ha impound-
ment of the Sabine River forms a 145 km border between Texas and Loui-
siana and is a popular bass fishing lake.

Native to a small region of southeastern Brazil (Forno 1983), S. molesta
has dominated water bodies over an expansive range including regions of
Africa, Asia, Australia and the South Pacific (Mitchell 1972; Forno & Harley
1979). Vegetative reproduction and rapid growth rates contribute to the
invasiveness of this plant, typically resulting in dense surface mats that cover
open water, degrade aquatic habitat, and constrain the use of reservoirs and
waterways (Mitchell & Tur 1975; Thomas & Room 1986). Until now, S.
molesta has been successfully intercepted at nurseries, botanical gardens, and
at a private pond (Myers 1982; Johnson 1995), precluding naturalization
in the United States. Likely sources for the introduction are local nurseries
found distributing the species as an ornamental water garden plant (Rhandy

elton, Texas Parks and Wildlife Department, personal communication).

This is the second Sa/vinia species introduced to these states. Sa/vinia
minima was first reported from Louisiana in 1980 (Landry 1981) and from
Texas in 1993 (Hatch 1995). Its distribution has increased extensively since
these reports (Montz 1989). The two species are readily distinguished by
the multicellular hairs on the upper frond surface. The apex of each hait is
divided into four branches. In S. molesta, the tips of the branches are joined
to form a cage-like structure, while in S. minima, branches are spreading
and free at the tips. Leaf hair features may be viewed with a 10X lens and
can be used by biologists for early field detection.

This report documents an introduction that constitutes a serious threat
to aquatic systems throughout the southern United States. Sa/vinia molesta
has been found above the high water level at public ramps and roads along
Toledo Bend Reservoir, likely dragged there by boat trailers, known vec-
tors for overland spread (Miller and Wilson 1989).

Sipa 18(3): 927-928. 1999

928 Sipa 18(3)

Voucher specimens: U.S.A. TEXAS. Harris Co.: Houston, wetland demonstration pond
of Robert Browning Elementary School, forming dense surface cover, apiculate sporocarps
in an elongated cymose system, 4 May 1998, Ronald K “Tones (FLAS, PIHG, TAES). LOUI-
SIANA. Sabine Parish: Many, Toledo Bend ees abundant in coves and shallows,

apiculate sporocarps in an elongated cymose system, 30 Nov 1998, James M. Hyde (FLAS).
ACKNOWLEDGMENTS

The author gratefully acknowledges assistance in identification from Nancy
Coile, Wendy Forno, eas David Mitchell.

REFERENCES
Forno, [.W. 1983. Native distribution of the Sa/vinta anriculata complex and keys to spe-
cies identification. Aquat. Bot. 17:71-83.
Forno, I.W. and K.L.S. Hartey. 1979. The occurrence of Sal/vinia molesta in Brazil. Aquat.
Bor. 6:185—187.
Haren, $.L. 1995. Salvinia minima (Salviniaceae), new to Texas. Sida 16:595.

JOHNSON, B. 1995. Giant salvinia found in South Carolina. Aquatics 17:22.
Lanpry, G.P. 1981 eee minima new to Louisiana. Amer. Fern J. 68:95.
Miicer, I. La and C.G. — 989. Management of Se/vinia in the Northern Territory. J.

Aquat. Plant Manage. 27:40—46.
Mircneit, D.S. 1972 “The K. ariba weed: Sa/vinia molesta. Brit. Fern Gaz. 10:251—252.
Mit

, D.S. and N.M. Tur. 1975. The rate of growth of Sa/vinia ie (S. auriculata
auct. 0) in laboratory and naa conditions. J. Appl. Ecol it 2213
Moniz, G.N. 1989. Distril inia mintma in Louisiana. Proc. | Aneel Meeting

Aquatic Pant Control Research ae 4—17 November 1988, West Balm Beach, Florida,
Misc. Paper A-89-1, U.S. Army Corps of Engineers, Vicksburg, MS, pp. 312-316.
Myers, F.D. 1982. Memo to state and territory agricultural i officials. U.S. Dept.
of Agriculture, Plant Protection and Quarantine, DA #
THOMAS, - . and P.M. Room. 1986. Taxonomy and eel ae molesta. Nature 320:
58 1—

ORTHODON VS MOSLA (LAMIACEAE)

NEIL A. HARRIMAN

Biology Department
University of Wisconsin-Oshkosh
SON WI 54901, U.S.A

MAN @ EWU, edu

The generic name Orthodon Bentham & Oliver (Lamiaceae) has never gained
widespread acceptance. Mos/a (Bentham) Buchanan-Hamilton ex Maximowicz
has been used almost universally for this Asian genus of about 22 species,
several of them quite weedy. Indeed, Bentham himself (1876) adopted Mos/a
and cited his own generic name in synonymy, without comment.

The relevant data are these:

Orthodon R. Brown, Trans. Linn. Soc. London 12:578. 6 April—August
1819, Mu
ee Bentham & Oliver, J. Linn. Soc., Bot. 9:167. 12 October 1865,

ae.

ea: ea Buchanan- Hamilton ex Maximowicz, Bull. Acad. Imp.

Sci. Saint Pétersbourg, sér. 3, 20:456. 1875, based on Hedeoma Persoon sect.
Mosta Bentham, Labiat. Gen. ae 366. 1834.

Orthodon Bentham & Oliver is an pce pate homonym of Orthodon R.
Brown, easily discoverable from Index Nominum Genericorum (Plantarum). However,
before plant taxonomy was blessed inn such invaluable compilations, there
was no practical and reliable way to discover the existence of duplicated
generic names. This was what led Bau and Nayar (1969) to propose conser-
vation of Mos/a against Orthodon Bentham & Oliver; McVaugh (1970) re-
plied (as Secretary for the Committee on Spermatophyta) that the existence
of Orthodon R. Brown (Musci) made Orthodon Bentham & Oliver (Lamiaceae)
a later homonym—the proposal was therefore superfluous, and was with-
drawn by its authors.

There the matter rested. Since 1970, no proposal to conserve Orthodon
Bentham & Oliver against Orthodon R. Brown has been submitted to Taxon,
Dan Nicolson informs me (pers. comm., 4 December 1998). One expects
none would arise, because Mos/a is so widely used that to return to a con-
served Orthodon would be destabilizing to nomenclature.

However, Orthodon Bentham & Oliver has now resurfaced: Gleason, H.A.
and A. Cronquist (1991, p. 445) adopt it in place of Mos/a. Its “impropri-
ety” is not remarked upon in Hammond (1992).

The name cannot be used; I surmise (following the suggestion of Paul Fryxell)
that its appearance may stem from the entry in Airy Shaw (1973), wherein

Sipa 18(3): 929-930. 1999

930 Sipa 18(3)

Orthodon is accepted in generic status and Mos/a is reduced to synonymy,
the compiler having been unaware of the existence of the priorable Orthodon
R. Brown (Musc1).

Mosla is represented in the United States of America only by the weedy
annual Mos/a dianthera (Roxburgh) Maximowicz, t

—

le type species of the genus;
its known range is Georgia, Kentucky, New Jersey, North Carolina, and
Tennessee (data from www.mip.berkeley.edu/bonap/checklist_intro.html),

but it can be expected to spread much more widely. It would be most un-
fortunate if the illegitimate name Orthodon dianthera were to become weedy
in the literature.

REFERENCES

Airy Suaw, H.K. 1973. A dictionary of the flowering plants and ferns, 8th ed. Cambridge
acne Press.

BENTHAM, 1876. Mosla. In: G. Bentham & W.J. Hooker. Genera Plantarum 2:1182.

Bau, C.R. and M.P. ei Proposal to conserve ene pereHe name 7333. Mosla Buch.-
teen Mai, (1875) (LAMIACI g 2 Benth. & Oliver (1865). Taxon
18:596.

GLEASON, 7 ‘a a A. CRONQUIST. ree en ul of vascular plants of northeastern United
States and adjacent Canada, ed. 4S

HamMOND, H.D. 1992. fe eceicne me addictions for and errors found in the Manual of
vascular plants of northeastern United states and adjacent Canada, Second Edition, by Henry
A. Gleason (late) and Arthur Cronquist. Bull. Torrey Bot. Club 119:88-89.

McVaucu, R. 1970. ee of the Committee for Spermatophyta. Conservation of generic
names, XIII. Taxon 19:81

TRIOSTEUM (CAPRIFOLIACEAE) IN TEXAS

WALTER C. HOLMES
Department of Biology
Baylor University
Waco, TX 76798-7388, U.S.A.
KAY M. FLEMING

Texas Parks and Wildlife Department
Athens, TX 75751, U.S.A
RUTH AND RON LOPER
University of Texas at Tyler
Der, TX ID 107 aS ah
JASON R. SINGHURST

Wildlife Diversity Program
Texas Parks and Wildlife il
Austin, TX 78704,

Trrostewm is a genus of about six species distributed in eastern North America
and Asia. These are coarse, perennial, erect herbs with one to many stems
arising from a woody crown. Many authors recognize three species from North
America, T. angustifolium L., T. perfoliatum L., and T. aurantiacum Bickn. (Small
1933; Fernald 1950; Steyermark 1963; Radford et al. 1968; Strausbaugh
& Core 1978). Triosteum angustifolinm is characterized by narrowed leaf bases,
whereas T. perfoliatum and T. aurantiacum have connate-perfoliate leaf bases.
The latter two are segregated primarily on the nature and amount of pu-
bescence and glandular hairs. Although recognizing the two species, Voss
(1996) mentions that “pubescence characters are not as consistent as some
keys suggest ...” Brooks (1986) treats them as infraspecific taxa, maintain-
ing that “The havik typically used to distinguish the two varieties, especially
stem vestiture and leaf shape, intergrade with some degree of frequency in
both the GP {Great Plains} and extraregional material examined. This suggests
that separation of the two entities is dubious.”

Correll and Johnston (1970), as well as Hatch et al. (1990) and Jones et
al. (1997), include only Triosteum angustifolium within the flora of Texas. Therefore,
the collection cited below is the first report of T. perfoliatum (T. perfoliatum
var. perfoliatum sensu Brooks, 1986) in the state. The species is known from
MA and NY, west to MN, eastern NE and KS, northeastern OK, south to
northern GA, AL, MS and AR (Fernald 1950; Brooks 1986). There is also

Sipa 18(3): 931-933. 1999

932 Sipa 18(3)
one record from Natchitoches Parish in northwestern LA (Thieret 1969;
Thomas & Allen 1996).

The Anderson County location, located at the western limits of the Pineywoods,
is a mature hardwood-pine forest dominated by Pinus taeda L., P. echinata
Mill. (Pinaceae), Quercus stellata Wang., Q. falcata Michx. (Fagaceae), Acer
rubrum L. (Aceraceae), and Liguidambar styracifiua L. (Hamamelidaceae). Common
understory plants include RAynchosia latifolia Nutt. ex T.&G. (Leguminosae),
Polygala polygama L. (Polygalaceae), Oplismenus hirtellus (L.) Beauv., Chasmanthinm
sesstliflorum (Poir.) Yates (Gramineae), and E lephantopus carolinianus Raeusch.
(Compositae).

The leaves of the specimen are distinctly panduriform and have margins
that vary from entire to conspicuously crenate-sinuate. Small (1933), Fernald
(1950), Radford et al. (1968), and Brooks (1986) all report strictly entire
leaf margins for all species of Trzostewm in the United States with the excep-
tion of the bases. Theret 26855 (LAF, TEX!), the only Louisiana collection
of the species, has similar crenate-undulate margins on some of its leaves.

—

—

her specimen. Texas. Anderson Co.: 3 mi WNW of Elkhart at Ivy's Preserve, 9 May
a Sashes Fleming, Loper, & Loper 6832 (BAYLU).
The following key, based upon Texas material, may be used to distin-
guish between the two species of the genus now known from the state.
1. Flowers yellow, | per axil; bracts subtending flowers two, exceeding the length
of flowers or fruits; calyx long hirsute, glandular hairs few; bases of midstem
T. angustifolium

leaves narrowed.
. Flowers reddish to orange, (1—)2 or more per axil; bract subtending flowers

one, ca. one-half the length of flowers or fruits; calyx densely puberulent,
T. perfoliatum

glandular hairs numerous; bases of midstem leaves perfoliate. ...........

Triosteum angustifolium occurs from central CT, Long Island (NY), and
PA, west to MO, southeastern KS, eastern OK, and east TX, south to GA
and northern LA. Correll and Johnston (1970) give the distribution as east
Texas, specifically Smith and Cass counties, both within the Pineywoods
Vegetational Region. Hatch et al. (1990) list the species as occurring only
in the Pineywoods. However, T: angustifoium is now known from the north-
ern part of the Pineywoods and also from the northern part of the Post Oak
Savannah in Lamar County.

Specimens examined. TEXAS. Cass Co.: between Linden and Avinger, 15 Jul 1969,
Amerson & Watson s.n. a same location, 3 Aug 1969, Amerson s.n. (SMU); 1 mi W of
Linden, off Hwy 155, May 1970, Correll & Correll 38089 (TEX). Lamar Co.: 500—

1000 ft W of Co. Rd. ve from a point ca. 1.8 mi NW of its jot with U.S. Re. 271, ca.
2.9—3.0 air mi NNW of j jct. U.S. Re. 271 and FM 2648 N of Paris, Camp Maxey (Texas
National Guard), N 33° 49! 25", W 95° 32'55", 4 May 1994, Carr & Wolfe 13737 (TEX).
Smith Co.: Swan, 17 Sep 1902, Reverchon 3208 an J).

HOLMES ET AL., Triosteum in Texas 933
ACKNOWLEDGMENTS

We wish to thank the curators of the following herbarium for the loan of
specimens or other information that made this study possible: BRIT/SMU,
SHSU, TAMU, and TEX/L

REFERENCES

Brooks, R.E. 1986. Caprifoliaceae. In: Great Plains Flora Association. Flora of the Great
Plains. Keren of Kansas Press, Lawre

CorreELL, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas
Research aah Renner.

FERNALD, M. L. 1950. Gray’s —— of botany. American Book Co., New York, NY.

Harcu, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas.
Texas ag ultural Experiment on pub. MP-1655, Texas A&M University, College
Statio

Jones, S. D. , JK. Wiper, and P.M. Monrcomery. 1997. Vascular plants of Texas: A com-
ea ae c es list including synonymy, bibliography, and index. University of Texas

Pre

con 7 E “HL E. AuLes, and C.R. BELL. 1968. ea of the vascular flora of the Caro-
linas. Univers of North Carolina Press, Chapel

SMALL, J. K. 1933. Manual of the southeastern flora. —— of North Carolina Press,
Chapel Hill.

STEYERMARK, J.A. noe Flora of Missouri. lowa State University Press, Ames.

STRAUSBAUGH, P.D. and E.L. Core. 1978. Flora of West Virginia, 2nd ed. Seneca Books,
Grantsville, WV.

Turret, J.W. 1967. Twenty-five species of vascular plants new to Louisiana. Proc. Louisi-
ana Acad, 32:78-82.

Tuomas, R.D. and C.A. ALLEN. 1996. Atlas of the vascular flora of Louisiana. Vol. I, Di-
cotyledons. Acanthaceae-Euphorbiaceae. Louisiana Department of Wildlife and Fisher-
ies Natural Heritage Program, Baton Rouge.

Voss, E.G. 1977. 1996. Michigan flora, Part HI, Dicots (Pyrolaceae-Compositae), Bloomfield
Hills, Michigan: Cranbrook Inst. of Sci. Bull. 61 and University of Michigan Herbarium.

934 Sipa 18(3)
BOOK REVIEWS

Diccs, Grorce M., Jr., BARNEY L. Lipscoms, and Roserr J. O'KENNON 1999.
Shinners & Mahler’s Illustrated Flora of North Central Texas. (ISBN
1-889878-01-04), hbk). Botanical Research Institute of Texas (509
Pecan Street, Fort Worth, TX 76102-4060) and Center for Environ-
mental Studies (Austin College, Sherman, TX 75091). Sida, Botani-
cal Miscellany 16. xiv + 1626 pp. $89.95. 174 color photos, 2,300
line drawings, color maps.

It has been a good many years since a genuine, bonafide complete hard-copy flora has
yy &

appeared for Texas or any substantial part of Texas. Thus it is with some excitement that
we nue this new book. The reviewer of this new FLORA was ces by the heft (at
almost 10 pounds it builds up the deltoids etc.) and 1600 pages crammed with informa-

tion. Obviously one cannot read every word and examine every illustration ina few hours
of perusal. My notes are pretty effusive—with a lot of overused adjectives they'll rate a
goose-egg in any journalism class—but who cares, so here goes: INCREDIBLE, FANTAS-
TIC, WONDERFUL, BE cn tes WOW! To be more specific: COMPREHEN-
SIVE, ENCYCLOPEDIC, INNOVATIVE, With every ee illustrated—it’s a new de-

parture in Texas floras, a breath of ore air. More than any other flora of the region, it approaches

the oft-stated goals of floristic workers to combine professional soundness and complete-
ness with usefulness and accessibility for people of all levels of botanical expertise, and for
all sorts of consumers of botanical information. In short, this is a super-fine product of a
dedicated, skilled, industrious and deeply knowledgeable team of experts, who are to be
praised and es aes along with their contributors and sponsoring institutions and
publishet

Now among the gushing words a little rain must fall. Perfection has been closely ap-
red, not fully attained. In some hours of review, about 20 misspellings have been

proac
detected, all falling into the category of minor inconveniences, none resulting in serious
misunderstanding. One literature citation was repeated under two different authorships.
But even a nit-picker like me can find very little to carp about.

turally one can differ with the authors on some of their taxonomic judgments. (Whenever

10 taxonomists are present, 30 opinions exist on any given issue.) But no need to belabor
that here. The authors’ judgments are all defensible, even those that differ from mine.

The beauty of the volume itself, including the section on superb color photographs, bespeaks
much loving attention to the details of lay-out, typography, binding, etc., and will add to
its appeal to the general public, those elusive intelligent lay persons ce may spot the
book in Barnes and Nobles and be captivated for the first time by a meaty type of wild-
plant botany, as opposed to the pabulum in the “wildflower” books.

Praise belongs not only to the team that produced the work, but to those nameless technoids
who have given us electronic sending and processing of data, words and illustrations, and
electronic layout and setting of type and illustrations. This reviewer thinks back to the
almost totally unillustrated Texas flora that came out in 1970, Every word was written
longhand, typed manually, edited, re-typed, edited, re-typed, edited and retyped. Finally,
the huge typescript was taken personally (by Donovan Correll, on the train) to a vast print
shop in Wisconsin where it was set (clackety-clack, clackety-clack) in hot lead on old Linotype
machines. Then the galley proofs were mailed to this reviewer in Munich, corrected by
him and by Correll, then sent back to Wisconsin for re-setting. Then the work was printed

Sipa 18(3): 934. 1999

Book REVIEWS fe ®
in an initial run of 3,000 copies. The signatures were assembled by hand and sewn to-
gether and the books bound (about one in three was defective on the first go-round). How
blessed we are to have a brand-new publishing industry in the last few decades in which a
lot of the processes are streamlined and quality is so much better controllec

So let us praise and congratulate everybody involved in this effort, and pulsliete this
great work vigorously so that it can become a part of the collections of every institutional
library and many thousands of private individuals. And let us encourage these authors and
others to produce equally useful floristic accounts for the rest of Texas and surrounding
regions (as they say they already plan to do), and to produce not only hard-copy but CD-
ROM versions and cyt s subject to constant up-dating. And let us encourage even
more foundations to support this type of mor so es it may accelerate and flourish as a
background informational field on which cons ffort y proceed and so that public
awareness and enjoyment of our wild treasures may acccienite, And all the people said:

MEN.—Marshall Johnston.

Taytor, RICHARD B., JIMMy RUTLEDGE, and Jor G. Herrera. 1997. A Field
Guide to Common South Texas Shrubs. (ISBN 1-885-696-14-0,
pbk.) A “Learn about Texas” publication from Texas Parks & Wildlife
Press, Wildlife Division. Distributed by: University of Texas Press,
P.O. Box 7819, Austin, TX 78713-7819. $19.95. 106 pp, 116 color
photos, 9 illustrations, | color map, 5 3/4” x 8 1/4”.

An informative, easy-to-use plant identification field guide to 44 of the more common
woody plants and cacti oF South Texas. aus handy sel eulee for landowners, land man-

agers, con

ost common
species of woody wereeen found in the South Tees Plains, and includes ea on on
their value to livestock, wildlife and human

The South Texas Plains area is a oo region, roughly south and east of the line
from Del Rio to San ae to Rockport. Also called the Rio Grande Plains or south Texas
“brush country”

_-
ct

‘ompasses about 20.5 million acres, covering fifteen counties and
portions 5 of fourteen ee Over 281 species of woody plants and 32 species of cacti are
recognized in the south Texas ecological region. The vast majority of these plants are found
in the lower Rio Grande Valley. The 44 described in this book represent an estimated 75%
of the overall eee and cacti biomass of the south Texas ecological region, including the
Rio Grande Valley.

In the i and 1500s, this area was dominated by grasslands, but woody plants were
often present in thickets, upland areas, major drainage’s and river bottoms. Natural fires
helped maintain the region as a grassland or savannah, reducing woody plant densities.
With the migration of early settlers and their livestock in the 1800s, brush densities in-
creased due to overgrazing, along with other factors such as lack of natural fires, soil com-
paction and — droughts. This increase of brush density was eventually considered
detrimental by ranchers. Brush control and range reseeding began in the late 1930s and
early 1940s, sere by the a research by wildlife biologists indicated extensive brush
control was detrimental to wildli e. Whit te- tailed eee hunting was on the increase by the

1950s giving landowners an economic incenti providing quality habitats. Since then,

>

Sipa 18(3): 935. 1999

936 Sipa 183)

brush removal has been reduced or applied in a manner that is not detrimental to wildlife,
thus improving the habitat for deer and other wildlife species such as birds, reptiles, rab-
bits, ground squirrels, bobwhite quail and wild turkey.

ideuvifieation of these key food plants and knowing their nutritional value becomes an
important aspect of evaluating habitat, range condition and ecosystem health of the South
Texas Plains area.

Grouped into thorned and thornless categories, the plants are alphabetized by family
with common and scientific names given. Each plant is laid out in a two page spread with
often three color photos showing habit, flowers and fruit. Text includes the description
Crude
Protein Value chart for each plant. A summary chart is provided showing the percentage
of nutritional values of crude protein (CP), digestible protein (DP), and digestible dry matter
(DMD) for each season of the year. In addition, a bibliography, illustrated glossary and

—

and values of the plant to wildlife, cattle, goats, as well as humans and a nutritiona

ae are provided.

With the ever increasing need to conserve our resources, it would seem to me that many
Pe plants could be incorporated into the South Texas homeowner's landscape. Native
plants not only provide beauty and ease of care, they can provide a welcomed food source
to some of the most coveted visitors into our gardens ... butterflies and hummingbirds. —

Linny Heagy.

PAUL KENRICK AND Perer R. Crane. 1997. The Origin and Early Diver-
sification of Land Plants A Cladistic Study. ISBN 1-56098-729-
4, pbk; 1-56098-7 30-8, hbk). Smithsonian Institution Press, 470 LEnfant
Plaza, Suite 7100, Washington, DC 20560. 202-287-3738, ext. 343.
$27.50 pbk, $55.00 hbk. 441 Pp.

This book represents the first attempt to synthesize the classical approach to land plant
evolution with the modern cau hes offered by morphogenetic and molecular evidence
presented in a phylogenetic context. The book is divided into 7 chapters; followed by 5
sa Sar aa an exhaustive bibhogr anne taxonomic and subject indices
sack ground, a PE imer on phylogenetic systemat-

e first two chapters give a historical b
ics, and the aplication ar paleobotanical data to modern systematic studies. It is followed
erstanding of the kingdoms of organisms and the

by a cogent Summary of our Current unc
higher ranks containing pant tne subsequent three chapters cover the Embryobiota,
Pol la and Lycophytes, and Lycopsida. Each section deals

with thé er eins ‘of the en s concape systematics, phylogenetic questions and aims of
analysis, choice of taxa, character descriptions and coding, analysis, results and discussion.
This organization of data is the first of ics kind, and clearly points out the depth or dearth

of our current knowledge for each character, its origin, Soe evidence from the
fossil record, and systematic utility. For each analysis and result, step-by-step explanations

are clearly given, limits of available data are thoroughly discussed, and suggestions for future
work outlined. The purpose of the series, “...co publish innovative studies! in the fields of

comparative evolutionary biology, especially by authors willing to introduce new ideas or
challenge or expand views now accepted.” is truly fulfilled in these chapters. I can chink of

Stipa 18(3): 936. 1999

Book REVIEWS 937

no more cogent summary of what we know, what questions remain, and how we might
proceed to answer them.

The last eee ‘Perspectives on the early evolution of land plants” is the most illumi-
nating of all. In addition to the proposed classification of the Chlorobiota to the level of
infraclass, it offers a clear explanation of such concepts as the annotated Linnaean conven-
tions recommended by Wiley, with several additions that logically permit the systematic
positioning of fossil groups. Comparison of the phylogentic classification presented for the
land plants, together with the historical and modern processes underlying character analy-
ses, give all students of plant evolutionary biology a fundamental understanding of the
strengths or weaknesses of competing concepts. I found the discussion of microphyll evo-
lution in lycopods onan complete, in which the three competing hypotheses: reduc-

ion, enation and sterilization, are explained in detail, first in ontogenetic, then in taxic
homologous terms

The appendices are detailed and give us a picture of how reliable fossil evidence is for
key groups, followed by complete character and character state coding tables. The bibliog-
raphy is extensive, as are the taxonomic and subject indices. This book is a must as a foun-
dation for any major graduate course in plant comparative morphology or systematic and
evolutionary biology. I wish it existed when I was preparing for my doctoral compre-
hensive exam!—John J. Pipoly II

James MauseTH. 1998. Botany. An Introduction to Plant Biology, 2nd
Ed. [Multimedia Enhanced Edition}. ISBN 0-7637-0746-5, hbk;
ISBN 0-7637-0406-7, CD ROM). Jones and Bartlett Publishers, 40
Tall Pine Drive, Sudbury, MA 01776. $75.00. 794 Pp. text, 20 Pp.
glossary, 23 Pp. index.

MArSHALL SUNDBERG. 1995. Instructor’s Resource Manual to Accom-
pany Botany: An Introduction to Plant Biology, Second Edition/
ANN M. MICKLE AND JAMEs E. MICKLE. Test Bank and Bio Art. (ISBN
0-03-005892-9, pbk). Harcourt Brace & Company. No separate price
given. 387 Pp. text; 37 Pp. illustrations.

This textbook, CD-ROM, instructor’s manual, and eae web site (http://www.jbpub.com/
botanylinks) provides the most comprehensive set of modern, multimedia course materi-
als this reviewer has ever seen. It is truly ee ie

The textbook is constructed in one of the most logical, easiest-to-use ways imaginable.
The preface clearly states that “three topics are so important, so fundamental, that they
must permeate every aspect of an introductory botany textbook and should be mentioned
or alluded to on every page: they are evolution by natural selection, analysis of botanical
phenomena, and diversity of organisms and all their components

he book is divided in four parts: plant structure, plant shastoléay and development,
genetics and evolution, and ecology. The “part openers” section that introduces each part
summarizes the chapters belonging to each part, and — those themes relate to the other
respective parts. Within each of the 27 chapters, a “concepts” section comprises several

Sipa 18(3): 937. 1999

938 Sipa 18(3)

pages and provides an annotated outline of the subjects chat will follow. There are “plants
and people” boxes in each chapter, that serve to explain the way in which che information
in the chapter relates to our everyday lives. These ees s are surely an “autapomorphic char-
acter state.” In addition to the plants aed people boxes, there are 27 boxed readings scat-
tered throughout the text that represent ancillary reading related to the subject treated in
the chapter. The accompanying interactive CD-ROM is referenced in the a an and
eac

1 sector of the CD-ROM is associated with a chapter or part of the book. The text is
richly illustrated with high-quality, full color photographs; SEM, TEM or an photomi-
crographs; and finally, color or black-and-white original arework, illustrating significant
structures and concepts. Other features, including the marginal notes, summary, impor-

tant terms section, review tia and BotanyLinks.net questions combine to provide a
formidable set of learning to

n

What immediately set fae text apart from any other I have seen is the nesting of the
information. Depending on the level of the stuc

QU

ents taking the course, selected saa of
the introduction can be used or skipped, and the supplementary materials provided on the
CD-ROM can be required or used as a een The coverage of topics, coordination
among the text, CD-ROM and n ling movie shorts), tables, charts, boxes,
artwork and photographs sum to immerse the Student in a truly multimedia experience.

The text is written in an excremely clear and concise style that painlessly introduces com-

plicated material in a step-by-step fashion. The book has obviously been tested and re-
tested for ease-of-use, clarity of presentation, and thoroughness, based not only on feed-
back given to Mauseth in the years he has used this ever-evolving text, but through the
comments of the thirty reviewers he thanks in the preface. Each subject area is absolutely
current and ts meshed with those that preceded it and those that follow. The book is printed
on high-quality glossy paper with a sturdy cover and attractive binding.

acillary materials for the instructor include the Lystructor’s Resource Manual, the Test
Bank and Bio-Art, a set of 150 full-color overhead transparencies, the Plant Biology Tutor
CD-ROM, the Instructor's CD-ROM, and a Video Resource Library
Vhe Instructor's Resource Manual contains 27 chapters, corresponding to the text, with
detailed instructions for setting up a lab to correspond with the chapter treated in the text.
In addition, the Resource Manual contains two appendices, “T

» Keys” and “Field Trips.”
The keys are for common trees of either the northeastern or southeastern U. S., while the
field trips cover areas including the Pacific Northwest, the Desert Southwest, the South-
eastern Borst and the Northeastern Forests. Each of the field trip. sections includes spe-

cific jor parks chat show selected habitats, a material | preparation
section, and a reference section. The Plant Biology Tutor CD- ROM i is menu-driven, easy
to install, and a treasure chest of information closely linked to the principal text. I have

not seen the overhead transparencies or the Instructor’s CD-ROM, but I assume that they
are of the same excellent quality as the other components of this fantastic set of teaching
cools.

In summary, I can unhesitatingly recommend adoption of this book by any department
offering introductory plant biology. It covers the subject in an exhaustive manner, has its
information uniquely nested, uniquely presented among written text, fantastic photographs

and artwork, an interactive CD-ROM and web site, and materials that require very little if

-John J. Pipoly IIL.

any modification for use in any geographical area.

Sipa 18(3): 938. 1999

Book REVIEWS

Kart Nikias. The Evolutionary Biology of Plants. 1997. ISBN 0-226-
58083-0, pbk) The University of ey inca Press, 11030 South Lan-
gley Avenue, Chicago, IL 60628. $24.00. 449 Pp.

tated by the author in Chapter 2, “The concept that individuals can be grouped into
ecis and that populations can be grouped into discrete biological units called spe-

cies is a central tenet in biology.” Niklas is to be commended for a treatment of both mi-

cro- and macro- evolutionary theory that combines basic elements covered in the classic

works on biological and evolutionary species, (Carlquist, Cronquist, Dobzhansky, Erh

Gould, Grant, Stebbins, et al.) with those of the phylogenetic school, (Eldredge, Cracraft

Wiley, Mishler, et al.), ina refreshingly new, synthetic way. The book is divided into four

parts, including a comprehensive review of “evolutionary basics”, “life’s chronicles: the fossil

record,” adaptive walks: a hypothesis,” and “long-term trends.” The book is the first I have

seen that has a strong botanical morphogenetic and paleobotanic emphasis, as well as a

peal

ich,

presentation of the adaptive walk concept.
The first part, dominated by a discourse on population genetics, is aimed at the advanced
undergraduate or beginning graduate piel and is written in a clear, concise manner. I

found relatively few omissions, with the exception of a discussion of character displace-

ment vs. the Wallace Effect. Homeven the early introduction of phylogentics is to be ap-
plauded, as is the historical review of Sewall Wright’s and others’ work with fitness. His
mphasis on heterochrony are rare in textbooks

introduction to species and speciation, with

of this kind. While I would have liked to see more detail regarding morphogenetic abbre-
an and additions (sensu Takhtajan and later, Funk and Brooks), the relationship of
those concepts to that of the puctuated equilibrium hypothesis offer a balanced review.
The concepts of hybridization, polyploidy and introgression are dealt with in a particu-
larly straightforward manner

The second part deals with the origins and early events in plant evolution, concomitant
with the invasion of land and air. Using molecular evidence, paleobotanical evidence, and
morphogenetic evidence, Niklas strings together a picture of early land plant evolution
that brings the student through the basics of molecular, cellular, histological, vascular and
dern contexts. It presup-

jon

reproductive evolution, first in their historical, then in their mo
poses that the student possesses a firm grasp of the comparative morphology of land plants,
but it is, nonetheless, a synthesis that does not exist elsewhere

The third part of the book, dealing with adaptive aquatic and terrestrial
most unique because of its ye aspects. It is far from light reading, because it
subjects, including tues. physiological and
owever, it iS written

walks” is the

contains some of the most complicated o
morphogenetic principles rarely Pee in an introductory te
so that one logically progresses from the unicellular cto malecnne then the aquatic to
the terrestrial habitat, with extensi discussion of the physiological, ontogenetic and ana-
tomical changes ee to alee Hee transitions. The only oversight I detected is a
reference to present discussions of genetic spirals and contact parastichal

consequence of m
Sets s simultaneously when teaching phyllotaxy.

The final portion of the book covers long-term trends, including divergence vs. conv
gence, and tempos vs. events. In the section on divergence and convergence, homology —
analogy are clearly differentiated, with examples ranging from the classic pe gune
and modern morphogenetic. While most texts in this area are dull and boring, the many
illustrative examples, and the quotations from Ernst Mayr and Yogi Berra certainly mak
it entertaining as well as instructive. While the discussion contains more inductive than

Sipa 18(3): 939. 1999

940 Sipa 18(3)

deductive reasoning than the chapters that preceded it, the synthesis of evidence from pa-
leobotanical to modern examples informs the student of the historical bases for modern
opinions. The final chapter, including explanations of cpDNA, mtDNA and nDNA, rates
of genomic evolution, molecular clocks, species orogeny and demise, followed by the fossil
record of speciation and extinction, offers another unique synthesis of information normally
scattered between micro- and macro-evolutionary course texts.

In summary, this book is an excellent choice for any one-semester course in plant evolu-
tion. It is a bargain that should not be missed. With supplementary reprints from perti-
nent works in population genetics, morphogenesis, and phylogenetic theory, it could eas-
ily serve as the backbone of a two-semester graduate course in plant phylogeny. I heartily
recommend it co any university biol logy instructor or student of evolutionary biology.

—John J. Pipoly HL

Stipa 18(3): 940. 1999

Index to new names and new combinations in this issue

salen tas papilloss Kral & M. Strong, sp. nov. 842
Schltdl. var. warnockii (1M. Johnston) Henrickson, comb. nov, 788

Amaranthus eiccciuli pensis Henrickson, sp. nov, 800
Amaranthus texensis Henrickson, nothosp. nov. 791
Amaranthus x tucsonensis Henrickson, nothosp. nov. 804

ener J.H. Kirkr.. nom. nov. 817
sai eich riedelii (Tul.) J.H. Kirkr., comb. nov. 817
j Kral & M. Strong, sp. nov. 844
tylis eleocharoides Kral & M. Strong, sp. nov. 846
see latifolia Kral & M. Strong, sp. nov. 848
Bulbostylis lombardii Kral & M. Strong, sp. nov. 850
Bulbostylis pachypoda Kral & M. Strong, sp. nov. 852
ulbostylis scirpoides Kral & M. Strong, sp. nov. 855

,e4

B
Bulbostylis spectabilis Kral & M. Strong, sp. nov. 85

Commicarpus coctoris N.A. Harriman, sp. nov. 079
ommicarpus praetermissus N.A. Harriman, sp. nov. 681
Croizatia brevipetiolata (Secco) Dorr, comb. nov. 832

Emorya rinconensis Mayfield, sp. nov. 694

)

Hymenandra acutissima (Cuatrec.) Pipoly & Ricketson, comb. nov. 738
ieee ns aie Pipoly) Pipoly & Ricketson, comb. nov. 736
emsl.) Pipoly & Ricketson, comb. nov. 718

ay

Hymenandta cot (Lundell) Pipoly & Ricketson, comb. nov. 739

Hymenandra pittieri (Mez) Pipoly & Ricketson, comb.

ae rdida (Lundell) Pipoly & Ricketson, here nov. 722

Hymenandra squamata (Lundell) Pipoly & Ricketson, comb. nov. 725
enandra stenophylla (Donn, $m) Pipoly & Ricketson, a nov. 715

Hymenandra wilburiana (Lundell) Pipoly & Ricketson, comb. nov. 728

M lia conspicua 7.K. lovee sp. nov, 819
Mu earn jaime-hintonii P.M. Peterson & Valdés-Reyna, sp. nov. 686

Myrsine luae Ricketson & 7 eats sp. nov. 747

Nuphar sect. Astylus D. Padgett, sect. nov, 824

Nuphar advena (Ait.) Ait. f. subsp. orbiculata (Small) D. Padgett, comb. nov. 824

Nuphar advena (Ait.) Ait. f. subsp. ozarkana (Mill. & Standl.) D. Padgett, comb. nov. 825
Nuphar advena (Ait.) Ait. f. subsp. ulvacea (Mill. & Standl.) D. Padgett, comb. nov. 825
Nuphar pumila (Timm) DC subsp. oguraensis (Miki) D. Padgett, comb. et sta. noy. 825
Nuphar pumila (Timm) DC subsp. sinensis (Hand.-Mazz.) D. Padgett, comb. et sta, nov. 825

Packera subnuda (DC) D.K. Trock & T.M. Barkley, comb. nov. 635
Phaseolus albescens McVaugh ex R. Ramirez & A. Delgado, *P. nov. 638
Shans newberryi A. Gray var. yesicola Sivinski, var. nov. 67

a hedemarkii Takeuchi, sp. nov. 770
Prunus gideonii Takeuchi, sp. nov. 705
Psychotria howcroftii oe sp. nov. 767

Spiraea xhitchocockii W. Hess & N. Stoynoff, hybrida nov. 827

Tripogon sivarajanii Sunil, sp. nov. 809

Tripogon vellarianus Pradeep, sp. nov. 811

Villasenoria B.L. Clark, gen. nov. 632

Villasenoria orcuttii (Greenm.) B.L. Clark, comb. nov. 633

A new endemic = of Mentzelia sect. Bartonia (Loasaceae) from New Mexico
ie tas Kk. Todsei
819

Nomenclatural novelties in Nuphar (Nymphacaceae)
Donald |. Padgett

823

A new hybrid of Spiraea (Rosaceae) from Oregon
William J. Hess and Nick A. Stoynoff

827

A new combination in Croizatia (Euphorbiaceae)
LJ. Dorr

831

Eight novelties in Abildgaardia and Bulbostylis (Cyperaccae) from South America
Robert Kral and Mark T. Strong

837

The flora of Indian shell rings from coastal South Carolina to northern Florida
Richard Stalter, Maritess Leyva, and Dwight T. Kincaid

861

Noteworthy vascular plants from Arkansas
Eric Sundell, R. Dale Thomas, Carl Amason, Robert L. Stuckey, and John Logan
arr

Noteworthy records of Mississippi vascular plants
Bruce A. Sorrie and Steve W. Leonard

889

Documented cogs ioeeti 1999:1. Chromosome numbers and pollen diameter
variation in T

spe

Lee F. Greer and A. Michael Powell
909

Salvinia molesta (Salviniaceae), new to Texas and Louisiana
Colette C. Jacono
927

Orthodon vs Mosla (Lamiaceae)
Neil A. Harriman

Triosteum (Caprifoliaceac) in Tex
W ee Holmes, Kay M. Fleming, ih and Ron Loper, and Jason R. Singhurst
31

Book notices and reviews 636, 678, 692, 700, 808, 860, 876, 888, 926, 934

V > | ISSN 00306-1488

CONTENTS

New plants pote Crater Mt., Papua New Guinea, and an annotated checklist of the species
’ Takeuc
l

eel

Vascular plants on a gypsum outcrop in southern New Mexico: A listing, a new variety and

taxonomic ree in the Anulocaulis leiosolenus complex (Nyctaginaceae), and a new
variety of Men zelia humilis (1. oasaccae)

Richard ae ee and Thomas Wootten

O87

A new species of Portulaca (Portulacaceae)
James Henrickson and Thomas Rk. Van Devender

Review of early nomenclature in Euthamia (Asteraceae: Astereae)

Guy L. Nesom
1009

A new Bulbophyllum (Orchidaceae) species from the Crater Mountain area in Papua New Guinea
N.HS. Howcroft
1019

New combinations in the Melastomataceae from Hispaniola
Henri Alain Liogier
1025

ennisetum advena sp. nov. (Poaceae: Paniceae): A common ornamental grass throughout the
southern United States

Joseph K. Wipff and J.P. Veldkamp
1031

New combinations in Setaria (P Paniceae)
William E. Fox, _ and Bei L. Hatch
1037

The taxonomy of Cyperus virens and Cyperus drummondii (Cyperaceae) in the southeastern
United States

_ Richard Carter, David K. Alexander, Charles T. Bryson, and Andreas Lazari
1049 : :

A new variety of]
Jeffrey K. Donabue «
1083

The genus Myrsine (Myrsinaceae)in
Jon M. Ricketson and Jobn J. Pipoly CONTRIBUTIONS
1095 pare renee

: TO BOTANY
Additions to the genus Ardisia subgenus Gt
pl Pipoly Ill and Jon M. Ricketson

¢

VOLUME 18
NUMBER 4
DECEMBER, 1999

Notes on nein see imitations in ene
Guy L. N D). Noyes
1161

REFERENCE GUIDE / PG. 1317

CONTRIBUTIONS TO BOTANY

FOUNDED BY
LLOYD H. SHINNERS
1962
Wm. Ff. Mahler
Publisher 1971-1992?

Barney L. Lipscomb
Editor
Botanical Research Institute of Texas, Inc.
509 Pecan street
‘ort Worth, Texas 76102-4060, USA
817 332-4441 / 817 332-4112 FAX
Electronic mail: sida@brit.org

Home page at the URL: hetp://www.brit.org/sida/

—

John W. Thieret Prof. Dr. Felix Llamas
Contributing Spanish Editor

Dpto. de Botanica, Facultad de Biologia

Associate Lditor
Biological Sciences Dept.
Northern Kentucky University Universidad de Le6n

Highland Heights, Kentucky 41076, USA L'-24071 Leon, SPAIN

Guidelines for contributors are available upon request
and on our Sida home page as well as the last page of each issue,
Subscriptions for year 2000: $27. Individual, $50, USA Institutions, $60, Outside USA;

numbers issued twice a year

&

© SIDA, CONTRIBUTIONS TO BOTANY, Volume 18, Number 4, pages 941-1316.
Copyright 1999
Botanical Research Institute of Texas, Ine.
-rinted in the United States of America
ISSN 0036-1488

NEW PLANTS FROM CRATER MT., PAPUA NEW
GUINEA, AND AN ANNOTATED CHECKLIST OF
THE SPECIES

W. TAKEUCHI

Botanical Research Institute of Texas
clo Lae National Herbarium and
Papua New Guinea Forest Research Institute
P.O. Box 314, Lae, PAPUA NEW GUINEA
ABSTRACT
An exploratory survey of the Crater Mt. Wildlife Management Area (CMW MA) has revealed
the presence of at least 1,200 vascular plant taxa within the 2,700 sq. km reserve. Included
among the findings are several new species and a significant number ofd fc stn utioaa records.
Three novelties are formally described and illustrated. The new beehlerii,
Jasminum pipolyt, and Psychotria matambua?. An annotated cl heck i ae elllecdous) is also

provided

The botanical documentation for the CMWMA is one of the most substantial for any
locality in Papuasia. The preservation status and floristic richness of surveyed communi-
ties are very high, comparing favorably with the best sites in Papua New Guinea. Contin-
ued application of the [CAD (Integrated Conservation and Development) strategy is rec-

ommended for the subject area.

ABSTRACT (TRANSLITERATED JAPANESE)

| ] ] | if

Crater zan yaseiseibutu nitai cl iyori 1,200 ij il |
bunrui guntai no sonzai ga 2,700 heihou iSiometona) inai de kiroku shoumei sareta. Hakken
saretamono no nakani fukumareteirumono wa kazukazu no shinshu to soutou naru kazu

no mezurashi ishu shokubutsu to bunpukuiki kiroku dearu. Mittsu no mezurashii shok
wa seishiki ni kijyutsu oyobi zukai ni yori shimesareta. Shinshu wa G/lochidion beehlerit, es
aes soshite me matambuai dearu. Chushaku tuki no shushu butsu ichiranhyou
utai sare

C rater zan ane kanri chitai no shokubatuseacnecd punked wa Papuasian chihou

gunraku no hozonjotai

nitotte mottomo kuwashi monono hitotus de aru. Cl
oyovi shokubutussou no yutakasa wa soutou ni takaku, Papua New Guinea no sairyou na
basho ni masaru hododearu. Kadai to sareteiru chiiki ni taisuru tougouteki hogo oyobi
kaihatu doryoku wo zokkou surukoto ga shourie sareru.

INTRODUCTION
Papua New Guinea's (PNG) biotic environments are a focal point for numerous
specialist studies, in large part due to the extraordinary diversification and

endemism of its flora. In an effort to preserve this biota, conservation int-
tiatives in PNG have been recently defined and prioritized by a number of

Stipa 18(4): 941-986. 1999

942 Sipa 18(4)

multiagency evaluations, of which the most consequential are the Conser-
vation Needs Assessment Report (Beehler 1993), and the Papua New Guinea
Country Study on Biological Diversity (Sekhran & Miller 1995). The Cra-
ter Mt. Wildlife Management Area (CMWMA) is listed by these apprais-
als in their site portfolio of localities with critical floristic or biodiversity
significance (Johns 1993: 26; Sekhran & Miller 1995: 121). Despite its pristine
condition and high-value biotic estimates, little information on the CMW MA's
botanical resources had previously been available. A floristically-oriented
census was thus initiated in 1997 to acquire data for determining the di-
versity and population status of CMW MA plants.

SITE SUMMARY

The 2,700 sq. km territory of the CMW MA (Fig. 1) encompasses a mosaic
of floristic environments ranging from alluvial communities at 150 meters
elevation to montane cloud forest at 2,100 meters. Except for scattered and
localized seres resulting from subsistance agriculture, the forest formations
are primary. Life zones from the lowland, premontane, and montane con-
tours are fully represented within the project boundaries. Surveys conducted
to date have examined alluvial swamp forest, lowland everwet forest, premontane
foothill forest, and stunted montane forest. Within the premontane and montane
zones, there are various Se hic-edaphic subunits with distinctive floristic

MA

assemblages and taxonomically characteristic seral sequences. The C

also includes aquatic habitats represented by lakes, oxbows, springs, per-
manent streams, and waterfalls. The various elements collectively comprise
an unusually comprehensive habitat suite, all located within an adminis-
tratively cohesive conservation unit accessible only by air.

GENERAL DISCUSSION

The following account of plant resources in the CMWMaA is based on
intensive collections from a two-year period of exploration. Forests near the
villages of Haia and Maimafu served as principal venues for the botanical
survey.

In Appendix 1, plant taxa are referenced by the specimen voucher or other
source for the occurrence claim. Nearly all registers are supported by fertile
collections made in multiple sets for international and local distribution.
The Lae National Herbarium (LAE) is the repository for unicates, first sets,
and holotypes. Recipients of duplicate sets have been designated in accor-
dance with LAE’s pee protocols, on which Kew (K), Rijksherbarium
(L), and Harvard (A), are the primary receiving institutions. Residual sets
will be allocated by LAE under their auspices, in compliance with previous
agreements made with our PNG host organization.

OR ENGAU.

ae

ron Soy
MORESBY » X
aed
)

Fic. 1. Island of New Guinea. Arrow: Crater Mt. Wildlife Management Area.

THONAAY J

‘

vouincy MON Ende, “IPl JoIVID WOT sdurig

£¥6

944 Sipa 18(4)

Collections were field-preserved with 70-75% isopropyl! alcohol and trans-
ported in plastic wrap to facilities of the PNG Forest Research Institute
(PNGFRD) for subsequent processing. Taxa encountered only in sterile condition
or otherwise not obtainable, were enumerated as a sight record if the plant
was unequivocally known to the writer (viz. ‘SR’ in Appendix 1). When-
ever possible, such sightings are referenced against LAE collections made
by other botanists within the Purari drainage; primarily by the former Department
of Forests or by CSIRO botanists L.A. Craven and/or R. Schodde. Other

collector citations are provided in Appendix 1.

—_—

he survey's original intention of developing a baseline census along the
CMW MA's full elevational gradient remains out of reach. In view of the
2,700 sq. km area of the reserve, this objective is an ambitious one under
any scenario. Although the total species content of the CMWMA remains
conjectural, establishing an estimate is still attainable with respect to checklisting
the plants likely to be encountered by researchers working in the area. No
empirical measure has yet been developed for the total number of plant species
enclosed by Papuasian habitat gradients, whether or not assessed on verti-
cal or horizontal lines, though Johns (1993: 26) estimates 2,000+ taxa for
the Crater Mt. area. The Johns’ estimate appears reasonable and is probably
conservative. But if accurate, this should mean that the present itinerary
has yielded up to 60% coverage of the CMW MA's total taxonomic content.

Of the floristic environments examined thus far, the forests around Maimafu
have received the most complete attention. As the CMW MA's high-eleva-
tion antipode, the Maimafu area was expected a priori to have the lowest
richness and to be the easiest to achieve collections saturation. Mid-mon-
tane stations like Herowana are not included in this summary, but since
such sites are ecologically interposed between Haia and Maimafu, many of
its plants are probably already checklisted by existing collections. The con-
spicuous lacuna is the alluvial formation represented by Wabo and compa-
rable sites at the CMW MA's low elevations, where floristic documentation
is still generally inadequate. The largest potential advances toward a com-
prehensive understanding of the CMW MA's flora will be made through critical
examination of the lowland alluvial zone, notwithstanding the earlier work
done by LAE and CSIRO in the Purari drainage.

A salient aspect of general appraisals of the Papuasian flora is the recog-
nition of a remarkably diverse orchid component with more than 2,000 species
(Schlechter 1911-1914, Johns 1993) and the existence of a comparably rich
complement of ferns (Parris 1974). Orchidaceous and pteridophytic elements

should normally peak in montane forests like Maimafu, but this has not
been demonstrated by the survey because of the unfortunate circumstances
imposed by the el Nino disturbance. The CMW MA was severely impacted
by the recent drought, with considerable losses noted among epiphytic plants.

Takeucui, Plants from Crater Mt., Papua New Guinea 945

Enumeration of orchids, ferns, and other herbs was severely preempted. Because
these assemblages are typically among the richest groups in the Papuasian
montane zone, it is certain that the survey coverage was less comprehensive
than if a comparable evaluation had been conducted under the conditions
usually prevailing. In comparison, fruticose and phanerophytic plants were
less affected by drought and responded to the resumption of rainfall with
mass flowering and fruit production. The woody plants are thus better rep-
resented in the expedition gatherings.

The checklist in Appendix 1 incorporates results from earlier surveys in
the Purari basin, conducted during the 1970’s by CSIRO and LAE bota-
nists. The LAE gatherings were made primarily in the Wabo area by what
was then the Department of Forests, and were part of the LAE institutional
number series. The CSIRO sets originate mainly from the lower part of the
drainage bordering the CMW™MaA proper, from areas subject to the same
habitat and forest conditions as are present at Pio-Wabo. These latter col-
lections were more extensive than the LAE survey and are numbered under
L.A. Craven and R. Schodde’s personal number sequences. The cumulative
CMW MA checklist presently includes 167 vascular plant families, 592 genera,
and ca. 1,200 morphospecies, but it is obvious that an undetermined num-
ber of taxa remain unrecorded.

On a comparative percentage basis, the low elevation ecosystem at Cra-
ter Me. is still unexplored and likely to reward future investigators with a
disproportionate number of discoveries. It can be noted in this connection
that the Maimafu collections consist largely of taxa already well-documented
from the Highlands region. The montane affinity of the CMW MA 1s clearly
with the general Highlands flora. However from the status of the Central
Ranges as a floristic barrier, ic can be surmised that lowland forests of the
CMWMA should contain peculiarized and endemic elements, as has been
shown by the novelty value of the CSIRO collections from low elevations.

DESCRIPTIONS OF NEW SPECIES

EUPHORBIACEAE

Glochidion beehlerii Takeuchi, sp. nov. (Fig. 2). Typus: PAPUA NEW GUINEA.
Cuimpu (SIMBU) PRovINcE: Crater Mt. Wildlife Management Area, E of Haia village,
hill forest with some anthropogenic disturbance, 6° 43'S, 145° 00! E, 775 m, 15
Mar 1997 (fr), W. Takeuchi 11,796 (HoLtotyPE: LAE; tsorypes: A, K, L).

—

Arbor mediocris, circiter 15 m alta, habicu caulifloro, ramulis teretibus. Folia (matura)
fere ovata vel elliptica, usque 17.5 cm longa, 8 cm lata. Capsula breviter obovato-pyrifor-
mis, 10-15 mm longa, 14—18 mm lata.

Subcanopy tree, 15 m height. Stem unbuttressed, outer bark pale brown,
not furrowed, slash pinkish-red, wood dense, straw. Branchlets slender, weak,
terete, distally fractiflex, indumentum hirtellous, brunneous, harsh. Stzpules

946 Sipa 1L8(4)

ES;

aK / ¢ fr
SS

ay

Wray a

CAM (| By | Hi
nat aa
VSS? rr,

Fic. 2. Glochidion beehlerii Takeuchi, sp. nov. A. Branchlet. B. Fruit cluster on main stem.
Drawn from holotype by N.H.S. Howcroft.

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 947

linear-acuminate to subulate, caducous or persisting through several nodes,
typically 3.54.0 mm x 0.3-1.0 mm, hirtellous. Leaves alternate, distichous,
decrescent on the branch, the lower ones orbicular-subcordate, 3.5—5.0 cm
wide, acroscopically with the leaves ovate and ultimately elliptic, to 10.5—
17.5 cm X 5—8 cm; blades thin-coriaceous or papery, sub-bullate, bichromatic,
adaxially dark green and glabrescent, abaxial surfaces strikingly blue-green
glaucous and with densely hirtellous nervation, larger blades gradually acuminate,
margin entire, base equal and obtuse; venation pinnate, laterals 4—7 antrorsely
directed pairs, arcuate, closing with commissural loops or not, crossing nerves
subscalariform, major veins prominulous above, more raised beneath, reticulum
ventrally obscure or invisible, dorsally manifest, areoles pulverulent; peti-
ole 3-4 mm long, slightly swollen, articulated at base. Inflorescence not seen.
Infructescence cauline from base of trunk, arranged in congested hemispheri-
cal clusters on moundlike eruptions of the wood. Schizocarp obovoid, pyri-
form, or depressedly globose, solitary or several together on a short ramification,
mature fruits 10-15 mm x 14-18 mm, 8-9 ribbed, 1—2 mm stipitate or
not, exterior surfaces subappressedly puberulous, rather hyaline, soft pink
in vivo, summit broadly recessed and centrally pitted, style column deciduous,
not seen; fruiting pedicel concolorous with exocarp, 2-12 mm long, pu-
berulent; calyx persistent, 6-partite, sepals lanceolate, 2—3 mm X 0.5—1.25
mm, laxly hirtellous. Seeds 2 per cell, enveloped with a smooth orange arilloid.

Distribution and ecology.—G lochidion beehlerii is apparently a mid- or late-
seral species in forest regrowth. It is locally common at the type locality
but has thus far not been collected elsewhere. The ripe fruits are said to be
eaten by cassowaries.

Etymology.—The new binomial recognizes Dr. Bruce Beehler, a noted
ornithologist and frequent contributor to our knowledge of Papuasian natural
history. Dr. Beehler was formerly a senior ecologist for Conservation Inter-
national and is currently the Director for Environmental Conservation at
Counterpart International.

ParatyPe: PAPUA NEW GUINEA. Cuimsu (SimBu) PROVINCE: Crater Mt. Wildlife Manage-
ent Area, vicinity of Haia, near the Wara-oo streamcourse (first river E of Mt. Widau),
alluvial forest, 6° 43'S, 145° 00' E, 640 m, 30 Sep 1996 (fr), W. Takeuchi 11,303 (BRIT, LAE).
Glochidion beehlerii will not enter Airy Shaw’s (1980: 92—96) key because
of the cauliflory, a condition previously unknown for Papuasian representa-
tives of the genus. There are two cauliflorous species from western Malesia
(G. oxygonum Airy Shaw from the Moluccas, and G. cauliflorum Merr. of the
Philippines, cf. Airy Shaw 1969: 12), but the present taxon is incompat-
ible with either one.
The new G/ochidion produces large clusters of pyriform fruits at the base
of the trunk. Except for the fructification, it is superficially similar to G.
caloneurum in overall appearance.

948 SIDA 18(4)

OLEACEAE
Jasminum pipolyi Takeuchi, sp. nov. (Fig. 3). Typus: PAPUA NEW GUINEA.
BORDER OF CHIMBU (SIMBU) AND D GUL F PRovINCE: Crater Mt. Wildlife Management
Area, alluvial forest near Pio River, 6° 47'S, 145° 02' E, 440—460 m, 23 Mar 1997
(fl, fr), W. Takeuchi 11,866 pga LAE; isotypes: A, K, L).

1th f maximus , COCCO USqUue 35mm diametro.

Inter congeneribus Jasmino Papuasiac
Foliis trifoliolatis et glabris, inflorescentiis lepidotis.

Sprawling suffrutescent, or scandent vines. Svems terete, glabrous, glossy
green, inermous, virgate, without exudate. Leaves i it _exstipulate, epunctate,
entirely glabrous, pinnately trifoliolate, very ete ag toliolate but then
with an articulated petiole; rachis and petiolules green, adaxially sulcate;
leaflets induplicate, coriaceous, adaxial surfaces dark green nitid, abaxially
medium green, apex gradually acuminate, margin closely reflexed, pase cuneate
or rounded, lateral blades generally lanceolate, 6.5—9.0 cm x 1.75—3.0 cm,
terminal blades elliptic, 8-12 cm X 2.5—5.0 cm; venation pinnatiform, ‘midrib
channelled above, prominent beneath, laterals S—8, thin, plane or prominulous,
undersides with domatia-bearing axils, the domatia distinctly immersed,
crateriform, bearded, reticulum lax and irregular, dried leaves typically developing
characteristic indentations along nerves; petioles 12-23 mm long, concolorous
with stems, not grooved. Inflorescence terminal or from subapical axils, cy-
mose, monochasial or more often dichasial, obliquely ascending, the larger
ones paniculiform and leafy, to 17 cm length, all axes peltately squamulate,
bracts linear, to 6.5 mm long, persistent, granular, pedicels 4—7 mm long,
gradually thickened distally, weakly constricted at the calyx. F/owers (rehy-
drated measurements) bisexual, brevistylous, heterostyly not seen; calyx
funnelform, 2.0—3.0 mm long (exclusive of teeth) to 2.5 mm wide, slightly
accrescent, light green, tube glabrous, limb S—G denticulate, obscurely fimbriate,

denticulations linear-subulate, 0.2—1.0 mm long; corolla —— im-
bricate, white, entirely glabrous, obtuse in bud, BNE 10 mm X 2.0 mm,
costate, laxly tuberculate at first, lobes 5 sometimes 4, elliptic-oblong, 6—
8mm X 3—5 mm, mucronulate; androecitum elcid. stamens 2—3, ad-
nate to tube 2.5 mm from its base, filaments 0.5 mm long, anthers oblongoid,
3.5 mm long, connective inconspicuously apiculate; gynoecium glabrous,
superior, | mm height, ovary undulate at the summit, style 0.5 mm long,
weakly compressed and tapered, stigma punctiform, presented below the
filaments. Berry geminate (aberrantly with cocci 3 or 1), lobes rounded,
monospermous, 31—35 mm diameter, connate for ca. 20 mm, exocarp very
smooth, nitid, glabrous, hyaline, pearly white with suffused green mottling,
pericarp pulpy, greenish within. Seed white, hollowed.
Diffusely spread and locally frequent at the type
locality. Conspicuous in edge situations along streambanks and forest mar-
gins but also seen in otherwise closed forest environments.

istyibution and ology

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 949

3. Jasminum pipolyi Takeuchi, sp. nov. A. Habit of flowering plant. B. Dichasium. C.
ey: rom side. D. Flower, distal view of limb. E. Infructescence habit, with typical bi-
lobed fruit. F. Berry, ge trilocular form. G. Fruit in cross-section. Drawn from holo-
type by N.H.S. Howcr

950 Stipa 18(4)

Etymology.—The new binomial recognizes John J. Pipoly IL, a specialist
in Myrsinaceae and Clusiaceae. John Pipoly is a colleague and Head of Flo-
ras at the Botanical Research Institute of Texas.

Malesian_ Jasminum have been checklisted by Kiew (1994) with 9 species
accepted for Papuasia. Lingelsheim’s outdated key (1928: 18) is still the
only one extant.

Fruiting specimens of Jasminum pipolyi are immediately distinguished by
the exceptionally large fruits, which are strikingly translucent mottled. Also
characteristic is the combination of squamulate inflorescence and glabrous,
trifoliolate leaves which are crinkled along nerves when dried.

RUBIACEAE
Psychotria matambuait cree: sp. ae 4). Typus: PAPUA NEW GUINEA.
EASTERN HIGHLANDS PRO : Crater Wildlife Management Area, ridge com-

ylex above Abegarema village on the ie to Mt. Mopahveh, ee stature mon-
tane forest, near GPS coordinates 92802843 N, 28240 E, elev. 1,768 m, 02 Aug
1998 (4, fr), W Takeuchi 12,920 (HoLrorype: LAE; isotyee: BRIT).

] is ] ye ae a H ] ]

Psychotria howcroftii Takeuchi affine
non glabra, cum pedunculo breviori (non trichotoma ex ben.

Shrub or subarborescent in mature forest understories. Branchlets green,
fleshy, pithy, cylindrical and robust, collapsed in sicco, furfuraceous with
pale brown innovations at nodes, otherwise glabrescent. Stipules valvate, hyaline
or opaque, fugacious or semipersistent at subapical nodes, deltate, to 16
mm X 13mm, entire, weakly carinate, laxly provided with setuliform hairs
on dorsal and ventral surfaces. Leaves decussate, glabrous, spreading, whether
or not conferted; blades fleshy-coriaceous, lacking domatia, elliptic to oblanceolate,
more often broadest above the middle, 11.0—23.5 cm Xx 4.25—9.0 cm,
+concolorous, adaxially very dark green, abaxially medium green, bifacially
somewhat fuscous with drying, apex acuminate from an obtuse or tapered
summit, the acumen generally 1-2 cm long, base gradually narrowing toa
petiole 3-4 cm long; venation pinnatiform, major veins elevated, laterals
equispaced, 15—19 pairs, divergent, arcuate, abruptly turned near the mar-
gin but generally not closing, reticulum irregular, obscure or hardly raised
above, prominulous beneath. Inflorescence contracted, strictly terminal,
paniculiform, 2—4 cm long, ramifications verticelled or not from a peduncle
at most 1.5 cm long, ultimately cymose, caducously bracteate, rachises greenish,
provided with scalelike +crisped hairs, commissures furfuraceous. Flowers
(rehydrated measurements) seen in advanced bud only, cymes crowded, glabrous
without on all parts, pedicels proximally articulate; calyx turbinate, 7—8
mm X 6 mm, margin truncate or tearing irregularly, tube smooth, base
passing insensibly to the pedicel; corolla white, gamopetalous, obtuse in
bud, valvate, penta- or hexamerous, petals ligulate, 9-10 mm x 3.0—3.5

TAKEUCHI, Plants from Crater Mt., Papua New Guinea ow

D fis dt
f iy pega?

Fic. 4. Psychotria matambuat Takeuchi, sp. nov. A. Habit. B. Furfuraceous nodes on branchlet.
C. Stipule. D. Drupe, exterior aspect. E. Drupe in cross-section, with pyrenes and rumina-
tions. Drawn from holotype by N.H.S. Howcroft.

952 Sipa 18(4)

mm, tube cylindrical, 4-5 mm x 4—5 mm, throat marked with a 3 mm
ring of uniseriate-septate hairs, otherwise glabrous; stamens alternipetalous,
inserted in the sinuses, glabrous, filaments 2.5 mm long, anthers introrse,
basifixed, exserted, oblongoid, 3.0—-3.5 mm x | mm; style 8 mm long, but
?heterostylous, stigma 2-fid, 3 mm long, papillate, presented at the level
of the anthers; disk glabrous, shallowly excavated in the center. Drpes obo-
void, 19-21 mm x 14—16 mm (exclusive of calyx), orange-brown, peri-
carp at first entire, angulate and blackened after drying, apex crowned by
the erect calyx remnant, the calycine residue 4—5 mm long, tubular, cylin-
drical to infundibular. Pyrenes 2, equal, triquetrous, crested, endosperm filling
the entire cavity, ruminations transversal.

Distribution and ecology.—Psychotria matambuai 1s a montane umbrophile
restricted to understories of mature forest stands. It is known thus far from Mc.
Bosavi and the Crater Mt. area, being especially plentiful at the latter locality.

Etymology.—The new binomial recognizes Karol Matambuai Kisokau, a
former Secretary of the PNG Department of Environment and Conserva-
tion. Karol Kisokau had also served as the first General Manager of the Crater
Mt. WMA and is currently Director of Conservation and Research of the
Lae-based Village Development Trust.

—

Other specimens examined: PAPUA NEW GUINEA. Southern Highlands Province
Northern i of Mt. Bosavi, eee montane forest dominated by Syzyg/wm and pee
06° 28'S, 142° 50'E, elev. 1,800 m, 22 Aug 1986 (fr), O. G. Gideon in LAE 57,347 (BR
K,L,L i aces Hishiands ae Crater Me. 7 Idlife Management ee 7
above Maimafu airstrip to base of cascade at Mt.

Mopahveh, montane forest near GPS co-
ordinates 92802843 N, 28240 E, elev. 1,495—1,585 m, 16 Jul 1998 (A), W. Takeuchi 12,130
(A, K, L, LAE); Crater Mc. Wildlife Management Area, buttress ridge to Mt. Aretame above
Hauneabiibo village, natural growth montane forest, near GPS coordinates 9281348 N,

90 E, elev. 1,677—1,830 m, 25 Jul 1998 (f), W. Takeachi 12,602-B (BRIT, LAE).
The new species is allied to the recently described Psychotria howcroftii

Takeuchi of the Bismarck-Ramu foothills of northern PNG. The large drupes
with cyathiform calycine residue are characteristic of both taxa. Since P. matambuat
is apparently confined to the southern (Papuan) side of PNG, the species
pair provides yet another example of the evolutionary floristic separations
imposed by the Central Ranges.

Although similar to P. Aowcroftii in foliar, stipular, and fruit characters

P. matambuai is distinguishable by the furfuraceous nodes and a congested
inflorescence with an indumentum of setiform hairs. On Sohmer’s (1988)
revision of non-climbing Psychotria, the new species should key to couplet

60 (ibid: 19). It can then be incorporated into the decision train by intro-

ducing a third line into the otherwise dichotomous sequence:

—

a Fruit 4—5 mm in length

ol
62

uc LO—-12 mm in length

TAKEUCHI, Plants from Crater Mt., Papua New Guinea Bis,

Fruit 220 mm in length Psychotria matambuai
P. howcroftii

The sister species P. howcroftii was erroneously reported in the type de-
scription as keying to Sohmer’s couplet 36 (Takeuchi 1999). The proper

result should have been to fork 60 together with P. matambuai. The two
species can be differentiated with the following couplet:
Branchlets furfuraceous at nodes; inflorescence 7 4 cm overall length,
monopedunculate, rachises hairy, cymes congeste P. matambuai
Branchlets glabrous; inflorescence to 10 cm ae length, ms
rachises glabrous, cymes loose P. howcroftii

There is a minor complication at fork 57, since the leaves of P. matambuai
and P. howcroftii can have more than 18 lateral veins, being typically 15-19
penninerved. However the suggested outcome is more plausible than the
alternative branch on the Sohmer key.

DISTRIBUTIONAL RECORDS AND OTHER NOTEWORTHY COLLECTIONS

ADIANTACEAE

Coniogramme sp. nov., aff. macrophylla (B1.) Hieron.; coll. 72,197. Dif-
fers from Papuasian populations of Coniogramme macrophylla by the longer,
narrower pinnae and lax venation. The collection needs to be checked against
C. macrophylla populations from western Malesia, to ensure that the charac-
ter states are outside the range of variation for the species as a whole. The
CMWMA plants are potentially new if the differences can be maintained
against extra-Papuasian material.

ANACARDIACEAE

Rhus lenticellosa Laut.; coll. 7/,30/, det. PE Stevens & WT. An unde-
termined Haia collection was placed to genus by P.F. Stevens (pers. comm.
6/98) and was subsequently keyed to the given result.

Rhus lenticellosa is apparently a rare vining species, formerly known only
from the Sepik and Southern Highlands Provinces (Ding Hou 1978). Th
conspicuously cuspidate leaves are defining. Lae Herbarium previously had
no specimens determined to this binomial.

BEGONIACEAE

Symbegonia parvifolia Gibbs; coll. 17,707. Apparently endemic to western
New Guinea, with the few LAE sheets of this taxon all originating in Irian
Jaya. Barkley (1972) had listed the species’ range simply as ‘New Guinea.’
Possibly a record for the eastern side.

BURSERACEAE

Canarium sp. ?nov.; coll. //,886. What appears to be a new species of Canarinm
was encountered during the recent survey. The suspected novelty has a de-

954 Sipa 18(4)

ciduous phenology, 2.5 cm acuminate stipules, and large leaflets up to 43
cm long and 14
resulted in identification. However the plant cannot be made to key out on
the Flora Malesiana and does not conform to the description of any known

Papuasian Canarium.

CLUSIACEAE/GUTTIFERAE

Mammea papuana (Laut.) Kosterm.; coll. 11,905, det. P.F. Stevens. Two
collections of this species were previously known, both from E. Sepik Prov-
ince (Stevens 1974).

CONVOLVULACEAE

Erycibe carrii Hoogl.; coll. 12,619. Keys to this result on van Ooststroom
(1955). Apparently rare. Lae Herbarium has a single sheet of this distinctive

cm wide: unusual features which should ordinarily have

species.

Erycibe hollrungii Hoog1.; coll. 1/,762.Twigs with longitudinal cork lines
and the densely strigose sepals readily distinguish FE. hol/rangii from close
congeners like E. hellwigii.

Hoogland (195 3a) described the species on the basis of a collection from NE
New Guinea. In the van Ooststroom revision (1953), the taxon was charac-
terized as being known only from the type, though Hoogland himself later
found the species along the Puria River in Madang (Hoogland 195 3b). Since
then there has been nothing reported on this apparently rare plant. Lae Herbarium
has a single sheet, consisting of the Hoogland number from Madang.

Erycibe hollrungii is relatively common around Hata and many duplicates
were secured during the recent expedition. The CMWMaA collections are a
first record for Papua.

ELAEOCARPACEAE

Elaeocarpus lingualis Knuth, ‘sepikanus group; coll. 11,672. A member
of a complex including E. e/atus, E. sarcanthus, and E. sepikanus sensu stricto.
Coode (1981) shows how to separate the species with flowering material,
but acknowledges the taxonomy of this group is still problematic.

The species is known from only a small number of localities and had not
been definitely reported from the Papuan districts.

EUPHORBIACEAE

Aporosa lamellata Airy Shaw; coll. / 1,826. Large oblongish leaves and
persisting auriculiform stipules indicate a relationship to A. petzolaris but
the conspicuously ridged capsules are unique in the genus (Airy Shaw 1978).
The fruiting collection keys unequivocally to the given species and matches
the type description in Airy Shaw (ibid).

Aporosa lamellata was formerly regarded as endemic to West Irya (ibid)
although a recent determination (by A. Scl rot, 1996) on NGF 42,544 places

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 993

the taxon in the West Sepik as well. The species is now recorded for the
Papuan side.

Baccaurea papuana E.M. Bailey; colls. 12,563, 12,948. (Fig. 5). The Maimafu
collection from 1,768 m is the highest elevation of record for the species,
which ts ordinarily characteristic of lowland and coastal environments. CMWMA
montane populations have a number of nonconformist features, including
leaves smaller than the low elevation form and much larger fruits 35—40
mm long and 27-42 mm wide. The collections were initially prepared as
sp. nov., but Airy Shaw (1980) adopts a broad circumscription of B. papuana
and his conservative view is followed.

Macaranga reiteriana Pax & Hoffm.; colls. 12,262, 12,274, 12,507. (Fig. 6).
A subarborescent from Whitmore’s (1980) ‘Longistipulata group,’ Macaranga
reiteriana is distinguished by a dense sericeous indumentum and by solitary
sessile fruits inserted on bare peduncles.

The species is known only from Morobe Province, Gulf Province, and
the Idenburg River in West Iryan (ibid). Collections 12,262 and 12,274
key directly to this binomial and agree with cited material. The species was
also recently reported from the Bismarck Mts., Western Highlands (Takeuchi
1999), and is probably much more widespread than the number of collec-
tion localities would suggest.

FABACEAE/LEGUMINOSAE
Inocarpus sp. nov. (‘rubidus’ morphospecies fide Verdcourt); colls. 17,139,
12,054. This cryptic species has been discussed by Verdcourt (1979) and keys
out to the given result. We have several collections from various expeditions
to the Papuan districts, so the taxon is apparently not particularly rare.
An unusual feature is the red sap. Leaves are distichous and with con-
spicuous raised areolations on the underside. The rather small stipules are
another identification aid, being subpersistent and often conferted at the
apical bud.
Our several sheets of the undescribed taxon all dried to a distinctive or-
ange-brown color.

FLACOURTIACEAE

Ryparosa calotricha Mildbr.; coll. 12,008. The species is recognizable in
sterile condition, due to its distinctive foliar characteristics. Not previously
recorded from Papua (Sleumer 1954), and although other Papuan collec-
tions may have been obtained since Sleumer’s revision, the tree is probably
uncommon in the southern provinces.

MONIMIACEAE

Palmeria gracilis Perkins var. ?nov.; coll. 12,431. The collection is atypi-
cally strigose on leaves and infructescence. Possibly a new variety.

956 Stipa 18(A4)

Fic. 5. Baccaurea papuana FM. Bailey. A. nme habit. B. Infructescence on old wood.
Drawn from Takeuchi 12,948 by N.H.S. Howcro

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 957

Fic. 6. Macaranga reiteriana Pax & Hoffm. A. & B. Habit of fruiting branch. Drawn from
Takeuchi 10,496 by N.H.S. Howcroft.

Steganthera sp. ?nov., aff.‘myrmecophilous group;’ coll. 12,742. Branchlets
are nodose and ant-inhabited but the surfaces are manifestly muricate, im-
mediately distinguishing the plant from the related S. Aospitans. The large
leaves are also distantly toothed rather than entire. Female receptacles are
distinctively 3-lobed and crenate. Possibly a new species.
MYRSINACEAE
Loheria reiniana (Jacobs) Sleum.; coll. 11,921. There are two endemic species
of Loheria, both of which are rare and little-represented in herbaria.
Sleumer (1988) cites a total of 5 collections of this species. Most of the

958 Sipa 18(4)

existing specimens have originated from the Papuan districts, but the spe-
cles 1s apparently nowhere very common. Lae Herbarium has nearly all the
exsiccatae known to Sleumer, including an isotype. There is no published
information on the ecology of the Papuasian taxa.

Loheria reiniana is locally common near Haia, where it was seen growing
as a terrestrial monocaul with leaves crowded in a terminal coma. The spe-
cies is always found only in sciophytic situations sequestered within ma-
ture stands and is never in seral environments. Blossoms fall quickly after
opening, generally on the same day that the perianth deploys.

Population-based studies have never been made of Papuasian Loheria and
the CMW MA occurrences afford that opportunity.

MYRTACEAE

Syzygium sp. ?nov.; colls. 17,235, 11,719. The collections are from a very
distinctive species which does not key on Hartley and Perry (1973) on any
couplet permutation. The suspected novelty is a subcanopy monocaul with
flowers in condensed cauline cymes.

OLEACEAE

Chionanthus sp. ?nov.; aff. salicifoltus (Lingelsh.) Kiew; coll. 12.965. The
herbaceous leaves with closely set venation are unusual for chis genus. Similar
to Carr 14,121, annotated by Kiew as Chionanthus aft. salicifolins.
ORCHIDACEAE

Vanilla wariensis Schltr.; coll. 17,842, det. N.H.S. Howcroft. Vanz/la is
the basis for a local commercial industry which has thus far relied entirely
on alien species. Endemic Vanilla have potential value as breeding stock for
introducing genetic improvements into the current horticultural base (Howcroft,
pers. comm.)

Vanilla wariensis is a rare orchid with exceptionally large blades and was

not previously represented at Lae Herbarium by any specimens. There are no
extant populations known of this species except for the Haia colonies (ibid).
RUBIACEAE
Psychotria dieniensis Merr. & Perry; coll. 12,343. Previously known only
from the Fly and Sepik drainages (Sohmer 1988).
Psychotria polita Valeton, or aff.; coll. 12,397. Psychotria polita is known
from the single surviving collection cited by Valeton (1927). Sohmer (1988)
does not include the species in his key since the plant is not sufficiently
understood. However there is a plate of the only extant sheet and a brief
description of its salient characters (ibid).

The Maimafu voucher agrees with the available information on this bi-
nomial, including the particulars of its leaves and the extremely delicate

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 959

axes of the inflorescence. It is either the elusive P. po/ita, or a new taxon
allied to it,

Psychotria sphaerothyrsa Valeton; coll. 12,979. A rare species. Known
previously only from the lectotype and isolectotype from West Sepik Prov-
ince and from a citation by Merrill & Perry (1946) from Central Province
(Sohmer 1988).

Psychotria womersleyi Sohmer; colls. 12,028, 12,056. Previously known
from only two specimens originating in Eastern Highlands Province, above
1500 m (Sohmer 1988).

RUTACEAE
Wenzelia tenuifolia Swingle; coll. 12,958. The entire genus is rare; all 5
species being known from very few specimens. Stone (1985) had discussed
the difficulty of evaluating Wenze/ia due to the scarcity of collections avail-
able for study. He also commented on the elevational range of the genus,
noting that Swingle (1967) had not thought it possible for the aurantioid
taxa to ascend above 1,220 m.

he present collection from 1,770 m is the highest elevation of record
for Wenzelia and is the fifth specimen known.

SAPINDACEAE
Alectryon myrmecophilus Leenh.; coll. 12,746. Previously known only
from Morobe Province (Leenhouts 1994),

Harpullia camptoneura Radlk.; coll. 17,889. A species of NE New Guinea
(Leenhouts & Vente 1994). Newly recorded for the Papuan districts.

SOLANACEAE
Solanum dendropilosum Symon; coll. //,854, det. D.E. Symon. The Crater
Mt. number was originally distributed as ‘sp. ?nov. aff. dendropilosum’ but has
been more recently determined to the species by Symon (pers. comm. 7/98).
Solanum dendropilosum is represented by few collections and the atypical
features of the survey voucher probably represent juvenile characters. However
the indumentum is unusual and possibly cannot be accommodated under
the present name (ibid).

Solanum peranomalum Wernham; coll. 77,204, conf. D.E. Symon. Col-
lection 11,204 was keyed to this species, formerly known only by the type
specimen from Mt. Carstensz in West Iryan. The determination has been
confirmed by Symon (pers. comm. 7/98).

new record for Papua New Guinea.

STERCULIACEAE
Sterculia monticola Mildbr. var laxiflora Tantra; colls. (fr) 72,148, 12,647.

960 Stpa 18(4)

The collections key to the given result on available characters. Since the
fruits from this species were previously unknown (Tantra 1976), an acces-
sory description is provided from the present material:

Fructescence racemiform, axillary, pendulous, ebracteate, fruiting stalk
to 11 cm long and 4.5 mm thick, tomentulose, the hairs densely and appressedly
stellate, mostly 4-8 armed; pedicels 20 mm long, articulated above the middle,
indumentum like the rachis; follicles 1-4, free, shortly stipitate, divergent,
coriaceous, mucilaginous, oblong-acuminate, to 9.0 cm long and 2.0 cm
wide, arcuate along the dorsal suture, ventrally somewhat plane; valves 3—
5 mm thick, exterior surfaces at first brown, later orange, striate with dry-
ing, scabridulous, covered by minute stellate hairs, these with arms +erecto-
patent and setiform, interior surfaces glabrous; seeds 8—11, black.

APPENDIX |. LIST OF PLANT TAXA FROM CRATER MOUNTAIN

Voucher source for occurrence record: AM = Andy Mack, C&S = L.A.
Craven & R. Schodde, GW = George Weiblen, Ha = village collectors ex
Hata, He = village collectors ex Herowana, LAE = staff collections from
the Lae Herbarium number series, S&C = R. Schodde & L.A. Craven, WT
= W. Takeuchi; sn = sin numéro (without number); SR = sight record of
taxon known to the project botanist. Other collectors indicated by name.
Determinations by WT unless otherwise noted.

FERNS AND FERN ALLIES Asplenium scandens J. Smith ex _ WT
11861, 12157, 12167, 125
Aspleninm steerei Hein aren, va 12104,
12227; ‘cromwellianum’
Asplenium (close to) steerei Harrington, WT
2250, 12433, 12491, 12493
Aspleninm subemarginatim Rose nst., WT 5.2.

ADIANTACEAE
Contogramme sp. nov., aff. macrophylla (B1.)
Hieron., WT 12197
Syngramma hie Brause, WT 11168
Taenitis blechnoides (Willd.) Swartz, SR,

Maimat
ees Aspleninm tenerum Forst., WT 124 83
ASPLENIACEAE Aspleninm unilaterale Lam., WT 12166

Asplenium acrobryum Christ, WT 11852, Didymochlaena truncatula (Swartz) J. Smith,

121%, 12554 — WT 1217
\yplenium bipmmnatifidum Baker, WT 11816, Diplora sp., LAE 61104, Purari R.,

ri 2487, 1 2552 2 E ‘sweeten: = Phyllis longifolia (Pres)
A kas cuneatum Lam., SR, Hata; pos- O. Ktz

sibly A. affine Sw. Diplora sp., Roce rs Purari, det. Croft,

A. oar aie Kunze, WT 11962, 12741 = Phyllitis sp., aff. ?longifolia
Asplenium keysserianum eg WT 12910 aera enmR A

splenium morobense Copel., WT 12114

alee lee alia oe im.) Bory, SR, on

Aspleninm nidus L. var. a WT 11798
osenst., or aff., WT

: Le abe taide am., WT 11742
bes or >. malesicum

159
a, tum aff. bantamense group’, WT
LI2Z6: ALSO

Diplazinm cordifolium B\., SR, Haia

Tolecu ae WT
A be ch. ie ie Christ, WT
1723; possibly A. Reysserszanum Ros.

TAKEUCHI, Plants from Crater Mt.,

Diplazium schlechteri Hieron., WT 11794
AZOLLACEAE

Azolla pinnata R. Br., SR, Maimafu
BLECHNACEAE

Blechnum acutiusculum (v.A.v.R.) C. Chr.,

12354, 1280
Blechnum dentatum (Kuhn) Diels, WT 11734
Blechnum cf. deorso-lobatum Brause, WT

23:23
Blechnum orientale L., SR, Haia; cf. LAE
61165, above Purari R.
Stenochlaena areolaris (Harr.) Copel., SR
Haia

S tenochlaena mtlnet Underwood, SR, equiv.
to 13
Stenochlaena a (Burm. f.) Beddome,
a; cf. LAE 61111, Purari R.
CYATHEACEAE
yathea ar (Wall.) Copel.,

€ si iam (Sm.)J. Smith, WT’ 5.2.
Di R, upper Maimafu
DAVALLIACEAE

Davatlia solida Cae aes cf. CGS
783, Purari, det.

Davallia a =n WT 12703

Davallodes novoguineense es Copel.,
WT 12526, 12989

Humata dimorpha Copel., WT 11777

Humata pectinata (Sm.) Desv., SR, O-Pio

Humata tenuts Copel., WT 11780, 12223,
12614

Sephari cf. dorsalis Copel., WT 11949;
orsalis - sinusora grou

p
Scyphularia pentaphylla (B1.) Fée, LAE 61153,
Purari R.

NSTAEDTIACEAE
Dennstaedtia novoguineensis (Rosenst.) Copel.,

1822
Dennstaedtia scandens (B\.) Moore, WT 11744
Histiopteris incisa (Thunb.) J. Smith, WT
11685

Pteridium aquilinum (L.) Kuhn, WT 12799,
eae P. ae
DIPTERIDAC

Dipteris ae Reinwardt, WT 12232
Dipteris novo-guineensis Posthumus, SR,
Maimafu; less common than congener

Papua New Guinea 961

siaiipa mmr ee:
(Forst. f.) Tindale, WT

‘i is
Di Le ee phacosti a (Ces.) Copel.,
EL : iver; det. Grimes
ae ae (L.) s. lat.,
WT 1

EQUISETACEAE
Equisetum ramosissimum Dest. subsp. debile
(Vauch.) Hauke, WT 5.7.; O-Pio

GLEICHENIACEAE

inearis,
Ghichenia dupe Br Sk: upper! Maimafu
—— hirta Bl. var. candida (Rosenst.)
Holttum, WT 12320
Glen ae cf. sordida Copel., WT 12225,
subg. ita
GRAMMITID
a mmodon ae (Hooker) Moore, SR,

Dic ie ks linearis (Burm. f.) Underwood
2215

Bienes nine (Copel.) Copel., WT
849

12
Ctenopteris repandula Kunze, WT 11817
Crenopteris subsecundodissecta (Zoll.) Copel.,
2 , 1236

Crenopteris sp. ‘ ‘taxodioides -yoderi group’

SR, upp
Canons sp. 2 WT 7 2907

Grammitis sumatrana (Baker) Copel., WT
Grammitis SPs ie 28 129096
1.) Morton

WT 11964
Prosaptia alata (Bl.) Christ, WT 12224;
prob. syn. with Crenopteris
HYMENOPHYLLACEAE
Crepidomanes ss (Poiret) Copel.,
WT 1193
ines iphlebi Hides oe
Purari ee det. Cro
Trichon j (Pres ceenie. LAE
— 281, Wabo dam site
[vi v.d.B., LAE 66314,

LAE 61147,

EVOMIVEIIA

near es
LINDSAEA GROUP
Lindsaea cf. lobata Poiret, WT 12115,
possibly L. obtusa complex

962

Lindsaea lucida Bl., SR, Pio
pests microstegia Copel.,

788

WT 11737,
ee eet Smith, WT 11722, 12194,
12701

aie hella J. Smith) Mettenius ex
Var. a (Mettenius ex Kuhr 1)
ame 589
rs pulc te (Br rackenridge) Carruthers
Seemann, WT 12896
Lapliaes tenutfolia BL.,
dam site
Sphenomeris chinensis CL.) Maxon, SR,
Maimafu airstrip
Sphenomeris retusa (Cav.) Maxon, WT 12612
Tapeinidium longipinnulum (Cesati) C. Chr.
SR, Haia
peels novoguineense Kramer, WT
11824, 12344

LAE 66296, Wabo

Sec ce novoguineense Kramer, WT
12317

LOMARIOPSIDACEAE
Bolbitis heteroclita (Presl) Ching, WT 11819,
12409
Bolbitis — ie )Ching, WT 11154;
miniature for
Bolbitis rrvnlaris ae kenridge) Ching, WT
2569
Elaphoglossum novoguineense Rosenst., WT
LO

S

ier sintata C. Chr., WT 11755;
ale “Purari
es fas ere |.) Holttum, SR,
Haia; cf. S&C 4644
LYCOPODIACEAE
Huperzia att. ?carinata (Poiret) Trevisan,

a zia nummulariifolia (B\.) Jermy, SR,

Hagelin )Rothm., WT 12442;
S878, Purz
Huperzia ct. phlegmaria (L.) Rothm., WT
Huperzia squarrosa (Forse. f.) Trevisan, WT
12055

Huperza wa (near) sguarrosa (Forst. f.) Trevisan,

6309, Wabo dam sit
Lycopodium volubile Forst. £., W T 12811,

2851

Stipa 18(4)

Palhinhaea cernua (L.) Vasc. & Franco, SR,
Haia & Maimatu
MARATTIACEAE
Angiopteris evecta (Forst.) Hoffman, WT
2230
Marattia brassii Copel., LAE 61162
River, det. Leiden list
ia cf. novognineensis Rosenst., WT
11832

, Purart

care AE

/(Houtt.) Kuhn, WT
11790, 12435, "12496 12602, 12610,
12624, 12626, 1263

ei biserrata ne Schott, WT

we cf. falcata (Cav.) C. Chr., WT
111 - panei close to falcata

Nephro hii Christ, WT 12067
123

sae Baker, WT a rd

Oleandra werner? Rosenst., WT 16

OPHIOGLOSSACEAE
Ophioglossum pendulum L., SR: cf. CES 796,

Purari
OSMUNDACEAE
Leptopteris alpina (Baker) C. Chr., WT 12363
‘typical form’ sensu Johns
POLY PODIACEAE
A glaomorpha drynarioides (Hook.) Roos, WT
11147

A glaomorpha heraclea (Kunze) Copel., SR,
aimafu
morphe hierony
"] 252 co ea Oo ye,
lags cf. not i aad . ae C.
fe Pee!
babi Mamata Ee) C Saoal 4 var. mucronata,
11845, 12467, 12783
Drynaria sparsisora (Desvaux) T. Moore, SR,

22 (Brause) Copel., WT

pay a » 4Li7L,

Goniophlebinm persicifolinm (Desvaux)
eddome, WT’ 117119
Goniophlebinm subauriculatum (Bl.) Presl,
12313, 12501, 12512
Lecanoptert is mirabilis (C. Chr.) Copel., WT

lens (BI.) Donk, AM 403:

TAKEUCHI, Plants from Crater Mt.,

Leptochilus decurrens B\., WT 11963
Microsorum glossophyllum (Copel.) Copel.,
Microsorum linguiforme (Mettenius) Copel.,
ie , Purari River, det. Grimes
Microsorum papuanum Baker) Parris, WT
12457 68, 12631, 12805, =

Phymatosorus sp.
Macrosorum punctatum (L.) Copel., WT 11953
Microsorum rampans (Baker) Parris, WT

Microsorum ae (Christ) Copel.,
Phy MaltOSOrUS SP.
Phymatosorus sai (BL.) Pichi Sermolli,
W7 1881, 12025, 12426
Pier nigrescens (B1.) Pichi Sermolli,
WT 12726

Ph il cf. nigrescens (BI.) Pichi Sermolli,
VE 127630

Pyrrosia foveolata (Alston) Morton var.
foveolata, WT 11946
Pyrrosia foveolata (Alston) Morton var.
lauterbachii (Chir.) Hovenkamp, LAE
66321, det. Leiden list
Pyrrosia princeps (Mettenius) Morton, WT
1931

Jets

Selliguea albidosquamata (B1\.) Parris, WT
1792, 12613

Sellignea enervis (Cav.) Ching; sensu

rele WT 12151, 12832, ‘grami-

neous’ for

ee enervis (Cav.) Ching; sensu

wenkamp, WT 12315, ‘typical’ form

oa

ise) Hovenkamp ora

Wises (Br al
WT 11781
PSILOTACEAE
Psilotum complanatum Swartz, SR, Hata
Psilotum nudum (L.) Beauv., SR, Hata

PTERIDACEAE
Acrostichum speciosum WWilld.,

det. Leiden
Preris excelsa Gaud., WT 12377, 13002
Preris gardneri (Fée) Hooker, WT 12438
Pteris moluccana Blume, SR, Haia

Preris orientalis v.AV.R., WT 12608, 12730,

Preris pacifca Hieron., WT 11910
Preris schlechteri Brause, WT 12161
Preris tripartita Swartz, WT 11778

Papua New Guinea

963

Preris wallichiana Agardh, WT 13003
Preris wernert (Rosenst.) Holttcum, WT

SCHIZAEACEAE

PE aslo f.) Swartz, SR;
C&S 736, det. Croft
Ly ee salicifolium Presl, WT ye
Schizaea dichotoma (L.) Sm., SR; cf. SG
aes det. S&C
Schi ligitata (L.) Swartz, LAE 66355,

det eee list

SELAGINELLACEAE

ee GROUP

SEC 4481,

Selaginella ‘velutina-wallichii group’, WT
4; interspecific distinction not clear

ana (Cesati) C. Chr., SR, Haia
ain: (Alderw.) C. Chr., WT 5.2
— ieee (Willd.) Hol ttum, WT
243

Ta id “ heii (Willd. ) Holttum, WT
‘renata-vepanda group

THELY PTERIDACEAE

Amphineuron aft. immersum (Bl.) Holttum,

Chingia imponens (Ces.) Holttum, WT 11687
Plesioneuvon marattioides (Alston) Holttum,
WT 124

Be ceaee ris cf. eee rensis (Gepp) Holttum,
WT 1 s to P. rodigasiana

Sphaeros eee atas pi (Rosenst.) Holttum,
WT 11932

=f

S lees os heterocarpus (Bl.) Holttum,
WT 11793

Sphaer ee Inv ca ae f.) Holttum,
1122, Purari R., det. Leiden list
Sphaerostephanos mutiny ioe (Copel.)

a ttum, ie 918
nos Sal (Brause)

* iol et WT 1

Sphaerostephanos i aso: en (v.A.v.R.)
Honea WT 1182
Sf itis (1 Taisen SR, Haia

THY RSOPTERIDACEAE
Culcita sp.; straminea or villosa, SR, upper
Maimatu
VITTARIACEAE
Antrophyum alatum Brackenridge, WT
11933; LAE 61094, Purari, det. Croft

964

Antrophyum subfalcatum Brackenridge, LAE
66327, det. Croft

Vittaria elongata Swartz, SR, Haia
GYMNOSPERMS

ARAUCARIACEAE
Arancaria bunsteinitt K. Schum., SR, Maimafu

CUPRESSACEAE

Libocedrus papuana F.v.M. var. papuana, WT
12203

CYCADAC
Cycas scrati oe Fyv.M., LAE 61174, Purari
iver, det. K.D. Hill, also CGS 750
GNETACEAE
Gnetium gnemon L. or G. costatum K. Schum.,
V"]
ean mia BL., WT 12079
PODOCARPACEAE
Dacrycarpus ee (BI.) de Laubenfels,

WT s.n.
syste as D, Don, SR, Pio; cf.
7, det. de Laubenfels
Prunn in ys amara (BI.) de Laubenfels, WT
12304, 12827

DICOTS
ACANTHACEAE
Calophanoides angustata (Warb.) Brem., WT

¢

2265-A
Dicliptera papuana Warb., WT 12510
Graptophylum cf. pictum (L.) Griff., WT
11126

Hemigraphis reptans T. Anders., or aff., SR
of cor tb, Haia

Lepidagathis sp., es : og 3

Peristrophe sp., WT 1

pee is oe ae B. Hansen,

Ro klossii 8. Moore, WT 12072, 12890
ACTIN AE

Sauratia aff. ?calyptrata Laut., WT 12918

Raneseeanentinr soit C, Simich, WT 12017

oe cf. dufaurii (F. v. M.) Bailey, WT

2107-

es naumannu Diels, or aff., WT 11124,
1689, 12200, 12475, 11124, 11689,
distr. as aff. conferta

Stpa 18(4)

Saurauia schumanniana Diels, s.l.,GW 716B;
"11811, 12022
Saurania oo Diels or aff., WT
ii me

s ?Obtectae, WT 12
a 12690, on yee -
Sauvaiia e series Obvallatae, aff ?conferta
Warb., WT 12042, 12078, not the sp.
Sauraitia sp., series Ramiflorae, WT 12476
Saurania sp., series Setosae, WT 11810
Saurauia sp. A, unplaced aff., WT 12013,

Saurauia ie
12302

Saurama sp. B, unplaced aff., WT 12097,
12916
sae G Cael aff., Sands et al.
482, Put
AMARANTHACEAE
Achyranthes aspera L., SR, O-Pio
Achyranthes bidentata ne , WT 12576
Celosia argentea L., 1967

Trestne herbstit Haale . a 12801

ANACARDIACEAE
Buchanania arborescens (B1.) Bl., AM 336,
559; WT 12091
Bun ees macrocas sas Laut., SR, Haia &
cf. C&S 765, det. Ding Hor
C See permet br eee Volkens, SR, Hata
Dracontomelon dao (Blanco) Merr. & Rolfe,
AM 252
— bear oe te SEC 4492, E.
Phan nium timoriense Te ) Leenh., WT
2092
a lenticellosa Laut., WT 11301, det.
svens & ee as indet.
Rhus tattensis ern WT 12404
WT 11243
WT 12543,

Semecarpus aruensis Engl.,
eid bracteatus Laut.,
12

oa magnificus KK. Schum., WT 12678

Semecarpus papuanus Laut., WT 12048,
12063, 12065, 12693 (fr), 12749 (fr)

Semecarpus rostratus Valeton, LAE 61120,
Purari R., dec. Ding Hou

Semecarpus schlechteri Laut., WT 11828

NNON E
Cananga odorata Hook. f. & Thoms., SR;
ct. SGC 4499, E. Purari, det. S&C

TAKEUCHI, Plants from Crater Mt.,

Cyathocalyx papuanus Diels, SR; cf. SEC
4230, 4385, det. Leiden

Gontothalamus aruensis Scheffer, WT 11234

Gontothalamus imbricatus Scheffer, WT
71206, 11800

Haplostichanthus longirostris (Scheffer) van

e
Polyalthia ‘glauca-discolor group’, WT 12094
Polyalthia aft. ?’oblongifolia group’, WT

11160, 11215, 11913, distinctive; lat-
erals acroscopic, leaves areolate
Popowia cf. pisocarpa (B\.) Endl., WT 11248
Popowia aff. ‘pisocarpa group’, WT 11901
Pseuduvaria cf. grandifolia (Warb.) J. Sinclair,
WT 12024

Xylopia malayana (Hook. f.) & Thoms., or
papuana ra WT 11994
APOCY NACE
Alstonia ae ee Wall., ex G. Don, WT

1264
Alstonia scholaris (L.) R. Br., WT 12071
Alyxia markgrafii Tsiang, WT 11740
Alyxta a SP.5 cf. series Floribundae, WT 12300
Anode blongifolium Hemsl., SR, Pio;
C 4380, dee Middleton

Pe ay K. Schum., AM 1; WT

abe ae
Fin bovata Wallich, CGS 813, E.

it t. Fors
oo Pon L) W.T. Aiton, WT
een ternatensis Valeton, CGS 825,

Psa actus ee a Markgraf, WT
12468

Melodinus forbesti see WT 11710; LAE
6

ae eae : Moore) Fosb. &
Si 997

es sane es .) Markgraf, SR,
Pio; cf. LAE 66354, det. Middleto
ee ne Markgraf, WT 12309 fy)
251

ne eae Schum.) D.J.
Middleton, SR, O-P

Parsonsta sanguinea CWernharn) Markgraf
var. brassit (Markgraf) D.J. Middleton,
WT 12387, 12880

Papua New Guinea 965

to) sanguinea var. brassit,

Dp : ]
WT 11760

Parsonsia warenensis Kanehira & Hatusima,
WT 12731-A

AQUIFOLIACEAE
cf. Ilex, WT 12634
ARALIAC
Gastonia sane (Harms) Philipson, SR,

aia
Harmsiopanax harmsit &. Scham., WT 12205
Mackinlaya celebica (Harms) Philipson, WT

oo ee (Harms) Philipson,
298, 72
pele boer, a (Warb.) Philipson, WT

Osmoxylon micranthum (Harms) Philipson,

Osmiox ‘ylon Bat: (Scheffer) Becc., SR;
789, Purari, det. Bhilinson
Polyscias cf. ledermannii Harms, WT 11982
Schefflera schumanniana Warms. ssp.
schumanniana, WT 11130

ARISTOLOCHIACEAE
Parartstolochia momandul (K. Schum.) M.J.
Parsons, GW 743, 813; based on GW det.
ASCLEPIADACEAE
Asclepias physocarpa Schitr., WT 12796
Dischidia ovata Benth., SGC 4480, E. Purari,
det. Liddle
soa torricellensis (Schltr.) PI. Forster,
17 Tae det. Et ae
iii ecne., WT s.n.;
SEC 4705, det. PL. i
Hoya australis R. Br. Anew oe (K.D.
Hill) PI. Forster & con, WT 11867,
11947; det. PI. Forster
ee seo bachii K. as.
sd I. Forster
Hoya ies Schler., SGC 4482, Purari,
n, (not seen)

Hoya piestolepis Schltr., WT 11945; det. PI.
Forster

WT 11185,

Marsdeni luti R. Br.,
P.I. Forster

Marsdenia sp., cf. venusta P.I. Forster, cf.
CGS 700 (type coll.), det. P.I. Forster

WT 11748; det.

966

Sarcolobus globosus Wallich subsp. peregrinus
Blanco) Rintz, SGC 4461, E. Purari,
det. P.I. Forster
ani Laniense os rltr. ") P.L. Forster, WT
843; dec. PI.
oa. oblongus Rinez, cited by Rintz
(1980) I

from Purari
sl lee R. Br.,
Pura . PI. For

SEC 4490, E.
ASTERACEAE (COMPOSITAE)
Adenostemma lavenia (L.) Kuntze, SR, Haia;
also SGC 4500
Adenostenma macrophylum (Bl.) DC., WT
2024, 12062
Ageratum conyzoides L., WT 12797
Bidens pilosa L. var. minor (BL.) Sherff, WT
12006
ee clei ven ales i223
R, Maimatu
WT 11743, 12245

Blumea riparia (BI) DC.,

Blumea Lene (BL) DC., WT 12271,
2848, 12981

Cosmos pipes tus HBK, WT 5.7

Crassocephalum crepidiotdes en )S. Moore,

WT 12605, 12640

Emilia sonchifolia (L.) DC. var. aa
aaa Pn ie T 12795-

Wolf) DC., cy Haia
WT 12604

: ae ony ie L.f. ex L., WT 12878

oe cn. WT 129351

Micr as Sa pyrifoly ¢(Lamk.) O. Ktze, WT
1208 9

seit cordata (Burm. f.) B.-L. Rob., W7

630

Eaean eerie eee.

a. ?nov., WT 12322; vining, fulvous
lanate underleaf
Tagetes minuta L., SR, Hauneababo track
Lithonia diversifolia (Hemsl.) A. Gray, SR,
Haia & Maimafu
BALANOPHORACEAE
Balanophora papuana Schitr., WT
BALSAMINACEA
Impatiens hawkeri Bull, WT 12518
WT 12256:
or as Grey-Wilson’s ‘group 10

E2755

Impatiens lineavifolia Warb.,

BARRINGTONIACEAE
Barringtonta calyptrata (Miers) R. Br. ex
Bailey, cited in Payens (1967)

Sipa 18(4)

BEGONIACEAE
Begonia aft. 2angustae Irmscher, WT 11195,
1262 2S 2

Begonia pinnatifida Merr. & ee . r : : a

Begonia aft. pseudolateralis Warb.,
‘brachybotrys-psendolateralis’

Begonia serratipetala Irmscher, WT 11171

Begonia SP: ‘kaniensis es , WT 12082
1245 S, 8, det. Gid

Begonia sp., oo. group’, WT 12080,
2852

Begonia sp. A, unknown aff., LAE 61187,
above Purari R.

Begonia sp. B, unknown aft., WT 12030,

2142-A
es Sie ie seas Pee WT

Syn crams ae Wr 12178,
21, 42 395

Sy ae a Mert. & Perry, WT
12176

Symbevonia parvifolia Gibbs, WT 11701
BIGNONIACEAE
as one i (L.f.) K. Schum.,
CéS urari, det. van Steen
ree ae ona ) eerie ssp.
pandorana, 460
1 mane dbp )K.Schum., ae i
2687, 12821, 12
BIXACEAE
Bixa ovellana L., SR, Hata
BOMBACACEAE
Camptostemon schultzti Mast., CGS 799, Purari
BORAGINACEAE
Cordta subcordata Lam., CGS 872, Purari,
c ollectors
Cynoglossum hellwigit Brand, WT 12255

Tournefortia minutiflora Ried|, cited in Ried]
996)
Tournefortia sarmentosa Lamk., SR, Haia
Trigonotis inoblita F.v.M. var. inoblita, WT
12032, 12087, 12834

2087,

BURSERACEAE
Canarium acutifolium (DC.) Merr. var.
sr eal LAE 61090, Purart, det.

Canarium cf. asperum Benth 10
Canarinm maluense pane ssp. wee, SR;
C&S 699, SGC 4393, dets. Leiden

TAKEUCHI, Plants from Crater Mt.,

Canarium vitiense A. Gray, WT 11250,

11894, 12294, 12078

Canarinm sp. nov., WT 11886
CANNABIDACEAE

Cannabis sativa L., SR, cultivated
CAPPARACEAE

Capparis lanceolaris Lesch. ex Steud., GW

814, det. G

CAPRIFOLIACEAE
Sambucus canadensis L., WT 12074
CAR PTERIDACE
pie es ‘i , SR; cf. CGS 845,
Purari, det. Leiden
CARICACEAE
Carica papaya L., SR, ?naturalized Maimafu
area, also cult
sce nce ee
L.) Willdenow ex Roemer
& ‘Schu ie WT 1 ee oad
Stellaria media L., WT 1
CASUARINACEAE
Casuarina cf. oligodon L.A.S. Johnson, WT
12243

Gymnostoma papuana (S. Moore) L.A.S.
c T sn.

CECROPIACEAE
Poikilospermum inaequale Chew, AM 542:
ba pany, tee 12729
paxianum (Winkler) Merr.,

we 215; WT 11158

CELASTRACEAE
Celastrus monospermoides Loes., WT 12351,
12356

Cc se novoguineensis Merr. & Perry, WT
Zoo 1224, 1261 a
: ptt sp jue 120
rr ae ssp. moluccana
Bl.) Dine a WT 12775
Salacia sororia Mig., WT 12037
Siphonodon celastrinens Griff., SR, Pio; cf.
GG 4393,

od

CHLORANTHACEAE
Ascarina philippinensis C.B. Robinson, WT s.n.
Ascarina subsessilis Verdcourt, SR, Maimafu:
rom sessile leaves
Chlovanthus erectus (Buch.-
WT 12004

Papua New Guinea

Ham.) Verdcourt,

CHRYSOBALANACEAE
Maranthes corymbosa B1., iat
Parastemon versteeghit en & se SR, Pio
Parinart papuana C.T. White, WT 11890

CLUSIACEAE (GUTTIFERAE)

Calophyllum * eee complex’ sensu
Stevens fA L287
Catloph hylan Ppnonon Laut ee 254
Calophyllum s
ee oe osum eae ae O22,
Purari,
Garcinia pe re A.C. Smith,

WT

Garcinia celebica L., WT 11906; SGC 4218,
det. Ste

kia a fr eae WT 12561, 12866,

1294
a. dulcis Roxb.) Kurz, WT 11981,
12084
Sui role Stevens, AM 2 eee
ut., WT 11973, 12041
wr 11797

Garcinia aff. Lae Laut.,
Garcinia aff. 2jaweri WT 12047
Garcinia ct. klinkii Laut., WT 1 soe
Garcinia latissima ve An 40; S&C 4

E. Purari; WT 1
G. raveinia ae Laue WT 11
P25 1 a

il 1887
Garcinia mammeoides Kosterm., WT 11247
wr 12618

, WT 12282,

Garcinia warrentt E.v.M.,
Garcinia sp. A, unplaced af
12964
Garcinia sp. B, unplaced aff., WT 12651,
ee caw (Laut.) Kosterm., WT
, det. Stevens
COMBRETACE
Quisquatis in a L., WT 11996
Terminalia ee K. Schum., AM 240
Terminalia tmpediens Coode, SR; cf. SEC
4310, det. Coode
Terminalia kaernbachii Warb., WT 11983,
12016

Terminatia cf. macadamii Exell, WT 12037
‘Terminalis microcarpa Decne subsp. microcarpa,
9, E. Purari, det. Coode
CONVOLVULACEAE

Erycibe carrit Hoogl., WT 12619

968

Erycibe hollrungii Hoog)., WT 11762

Ipomoea batatas (L.) Lamk, SR, cultivated

Merremia sp., (probably) pe/tata (L.) Merr.,
SR, Hata-Pio

CORY NOCARPACEAE
Corynocarpus cribbianus (F.M. Bailey) L.S.
Smith, SR, Pio; cf. SGC 4452, Purari,
der. Molloy
CUCURBITACEAE
Cucumis sativus L., SR, cultivated
Luffa Lege (L.) Roemer, CGS 843,
urari, det. Frodin
oS aeek mucronata (B\.) Mig., WT 11813
CUNONIACEAE
Acsmithia sp., SR, upper Maimatu
Caldcluv. ‘aelebicr (BL.) Hoogl., WT 12214
— ee la ee (K. Schum.) Hoogl.,

ila sehles var. glabra, WT 12257;
typical form

Pullea ake Schltr. var. ais WE 12770;

=>
=

er f. ‘pers
pr aieis en i oe ve 12813
Schizomera (probably) parvifolia Perry, SR,
upper sins A ae ste ale
SR, Maimafu, sterile

Wernmannia sp.,
DATISCACEAE

Octomeles sumatrana Miq., SR, Pio
DICHAPETALACEAE

Dichapetalum papuanum (Becc.) Boerl., WT
11878

i a seeds Leenh., WT 11849
ably) timoriense (DC.)
F SEC 4427, det. Leiden

cn alum (pr
Boerl.

DILLENIACEAE
Dillenia castaneifolia (Miq.) Diels, SR; cf.
S&C 4370, det. Hoogland
Dillenia Mlebin Diss WT 12310, 12867
Tetracera nordtiana F.v.M. var. ides
(Mart.) Hoogl., SR on sp., Haia-Pio;
SGC 4365 for var.

DIPTEROCARPAC
Vatica papuana cei SR, Pio
EBENACEAE
Diospyros ct. sees J.R. & G. Forst.) PS.
en, WT 11850, 12014, 12663,
12839

Stipa 18(4)
Diospyros ferrea (Willdenow) Bakh., s.1., WT

986
Diospyros hebecarpa A. Cunn. ex Benth., WT
12764

ELAEAGNAC
Elaeagnus ae Roxb. var. triflora, WT

ELAEOCARPACEAE
Aceratinm ledermannii Schltr., WT 11840,
12919

Aceratinm muelleranum Schitr., WT 12079
Aceratinm oppositifolium DC., WT 12099
ia oe easel Warb , WT 12098

Schlet ., SR, Pio;

f. SEC 4 43206, 4401, det
Seas ledermannit Schitr SR, Pio: ae

Pullen 6455 near Purari
— gine ae WT 11672,

Ela multsetn Schler., SGC 44406,
NGF 41138, Purari

i | oe

sas noubuysii Koorders, SGC 4463,
E. Purari R., det. Coode

Elaeocar pus polydactylis a hitr., WT 12622

(fi de's ‘group 4

oo (c losest to) schlechteranus A.C.

mi 2538, possible sp. nov.

a ee sepikanus Schltr., s.1.,

Elaeocarpus sericoloides A.C. Smith, ee
det. GW

E oe oe (Gaertn.) K. Schum.,

>
ie)
yy
ie)
Oo
aae

)

Blanes sp., ‘sepikanus complex’, WT
2842 (fr), can’t separate spp. with fruits

Ena sp. Pe icus-altisectus BrouP
WT 3942

r), can’t separate without

flowers
Sericolea micans Schltr., WT 12374
Sloanea cf. nymanti K. Schum., WT 12766,
sut leaf size atypica
Sloanea att. pulleniana pas = 669
Sloanea Fv.M., 2, det. GW

ERICAC
ee aft. anchorifera J.J.Sm., WT

»)
Dimorphanthera brevipes Schltr., WT 11721,
P.E. Stevens

TAKEUCHI, Plants from Crater Mt.,

Dimorphanthera dekockti J.J. Sm. var.
Sie (Sleumer) Sleumer, WT 12/84,
12403

Dimorphanthera cf. elegantissima K. Schum.
var. ees (Sleumer) Stevens, WT
12041, 12305, 12406

ras sa alae Schler., WT

2187, 2937, nO det. Siang

Di ph , pers. com,
PE. Stevens

Gaultherieae indet., ?Gaultheria, WT
12451, could be new

Rhododendron dielsianum Schltr. var.
dielsianum, WT 12252, ais 12845

Rhododendron macgregoriae

va
12420, a 12555,

macgregoriae, WT 12
26 12800
Rhododendron multinervium Sleumer, WT

12026
Rhododendron phaeochitum Fv.M., WT 12278
Rhododendron scabridibracteum Sleamer, WT

1231
Rhododendron zoellert Warb., WT 11239
Vaccinium acrobracteatum K. Schum., WT
12318

Vaccinium carneolum Sleamer, WT 12316
Vaccinium cyclopense J.J. Sun. f. cyclopense, WT
1238]

Vaccinium finisterrae Schler., WT 12378

EUPHORBIACEAE

Acalypha grandis Benth., AM 211
Acalypha hellwigii Warb., WT 11197,
12629, 12672, 12675
Antidesma olivaceum K. Schum., WT 12074
pis _ ee K. Schum.,
11715; no result on ay

en aw
Antidesma cf. sarcocarpum Airy Shaw, WT
Dye oe concinnum ges
Aporosa lamellata Airy Shaw, 1826
Aporosa petiolaris fe Shaw, WT 12001
Baccaurea papuana FM. Bailey, WT 12563,
12948; also LAE 61083, Purari River,
det. Dockrill
Bischofia gavanica Bl., SR, Pio; cf. SGC 4688,

cei ath tokbra: (BL.) Kurz, WT 13009
cernua (Poir.) Muell.-Arg., WT
Pee 12506

Papua New Guinea 969

Breynia vestita Warb., WT 12798

Claoxylon lutescens Pax & Hoftm., WT 12258,
12574-A, 12992

Claoxylon cf. microcarpum Airy Shaw, WT
11844; ‘ledermannii-microcarpum group’

Claoxylon sp., ‘Purpurascentia group’, AM
598

Codiaeum variegatum (L.) Bl. var. be
(Decne.) Muell.-Arg., WT 71

Drypetes aff. lastogynordes Pax & ee WT
12082, 12

Endospermum labios Schodde, WT 11803

Euphorbia plumerioides a ai ex Hassk.
var. acuminata J.J. WT 12702

E piers) Muell.-Arg.,
WT s.n.; NGF 41139, Purari R.

Galearia celebica Koorders var. celebica, SR;

C 4240, det. Leiden
G lochidion chlamydogyne Airy Shaw, or aff.,

uk

Glochidion delticola Airy Shaw, S&C 4504,
E. Purari R., det. Leiden lis

Gti fara Migq., or oe. WT

Glubiton mitt ere ae Shaw, AM 650
Gl

ovo-guineense K. Schum., AM
590; WT 12231, 12635
Glochidion perakense Hook. f. var. supra-

axillare (Benth.) Airy Shaw, rr 619
sialon = nov.; aff. calonenrum Airy Shaw,
ues 11796
M toides Fv.M., GW 755, 756,
det. GW
Ma gustifolia Laut. & K. Schum.,
WT 11238
Macaranga ae: Laut. & K. Schum.,
WT 12204, 12264

Macaranga we Whitmore, WT 12087

Macaranga cf. ducis Whitmore, WT 12088

wii fallacina Pax & Hoffm., WT
1934

ee papuana\J. J. Sm.) Pax & Hoftm.
var. glabristipulata Whitmore, AM 609

Macaranga ptm Pax & Hoffm., WT
12208 62

oe ee iglandulosa Warb., WT
LL D2

Macaranga reiteriana Pax & Hoffm., WT
12262, 12274, 125

970 Sipa 18(4)
oo cf. papuana Sleumer, WT 11805,
12699, posiy ae eee

Pens SPs aff. ?hoffmannii Perry, WT Casearia ripicola Sleamer, WT 12162
2558, 1D:

parr tanarius (L.) Muell.-Arg., GW
det. G

>. dubiae Whitmore Flacourtia ae ‘it & a WT 13011
Homatinm foetidum (Roxb.) Benth., WT s.n.
oo. ) Benth., SR, Pio;
, det. Leiden
Pain un a. “AM 356, 629; WT
ae 744

ee eee Airy Shaw, LAE
15, Purari R., det. Airy Shaw

es paniculatus aewiMiwell, -Arg., WT

Mallotus sp. nov.; § Mallotus, WT 12159,

2454 aes as Mildbr., WT 72008
Manihot esculenta Crantz, WT 12795-B Trichadenia philippinensis cee. WT sin.
Octospermum pleiogynum (Pax & Hoffm.) Airy Xylosma papuanum Gilg., WT 12685, det

Shaw, SR, Haia Damas
se lieg novo-guineensis (Warb.) K. GESNERIACEAE
713, 12259 Aeschynanthus spp.,. WT 11112, 11113,
Phy nea Muell.-Arg., WT 12247, 11266, 11961

12281, 12809 Cyrtandra bracteata Warb., WT 11162,
ce cf. ciccoides Muell.-Arg., WT 11299

1226 Cyrtandra cf. decurrens de Vriese, WT 11151

Phin clambordes (Fv.M.) Diels, WT Cyrtandra att. 2elata Schler., WT 11270
Cyrtandya fusco-vellea K.Schum., WT 11181,

a a flav iflorus (Laut. & - — ) 11196, 11273, 11703
1aw, pr beet, de pbelO7 Cyrtandra b Lo 75] pidissi7 la Schler., WT 11157
Poy rheophilus Airy sae SR, Haia Cyrtandra cf. ganowskyi Schler., WT s.7.
Phyllanthus (closest to) rubriflorus J.J.Sm., a sp. A, § Centrosiphon, W7
WT 11259 ll ee 163
Phyllanthus uvinaria L., WT 12676 cy - he oo WT
ee ‘on amboink vain eee , WT 11995 11 ae a 14
SR, naturi alized M aimafu ee sp. C, § Cenerosiphon, WT
area 11267, 112 11292, 12996
EUPOMATIACEAE Cyrtandra sp. D, § Centrosiphon or
Eupomatia lanrina R.Br. WT 11098, 12043 Loxanthe, WT 11289,
FAGACEAE Cyrtandra sp.E, § Diplochiton, WT / gus
ss seciie aes )A.DC.,SR, SS. sp.F, ae ere WA

Cyrtandra sp. c. § Geodesme, WT 12776
aig _ H, § Geodesme, WT 12027,
2492 >)

Onis ee (Miq.) Rehder, WT s.7.,
carpological coll.

Lithocarpus lauterbachii (von Seemen) r

Markeraf, wr 13 O10 coi sp. 1, ses alae ra, WT
Lithocarpus rufi (Markgraf) Rehder, 12052, 12138, 1234 ’ ;
Cyrtandra sp. J, (Cyrtandropsis), §

WT 11944, 12219, 12472, 12777
Lithocarpus schlechteri Markgraf, WT 123006
Lithocarpus ct. schlechteri Markgrat, WT

:

ra, WT 11282, 11960,

Leucocyrtanc
12854

Cyrtandra sp. K, eel Ae Cen)
eee eens WT

FLACOURTIACEAE tees Cyrtandra sp. L, § oe il m/
Casearta clutiacfolia BL., WT Rae 2 Phaeonicinn Wr 11970 83
Casearia cf. clutiaefolia Bl., WT 12042 Cyrtandra sp. M, § aceocoieanten, wr

Casearia grewiaefolia Vent., or atf., WT 12855

TAKEUCHI, Plants from Crater Mt.,
pea sp. N, § Prosthecisiphon, WT

cone _ O, § Rhabdocyrtandra, WT
1291, 11769
coment sp. P, unplaced, WT 72091,
1247
a sp. Q, unplaced, WT 11172
(filibracteata facies)
Cyrtandra sp. R, unplaced, WT 12092

Dichr le spp., or as Agalmyla, WT

, 11226, 11708, 11807
Rhy noch discolor (Maxim.) B.L. Burtt,
il eo)
GOODENIACEAE

Scaevola oppositifolia R. Br., WT 11138,
12593, 12983
GROSSULARIACEAE
Carpodetus arborens (Laut. & K. Schum.)
Schltr., AM 616
Polyosma att. amygdaloides Reedet
2073, 12386; keys to P. stenosiphon but

ot that sp.
Be ‘cestroides- induta group’, SR,
Maimafu
Polyosma forbesii Valeton, or aff., WT 11249,
184]
HALORAGACEAE
Gonocarpus halconensis (Merr.) Orchard, SR,
upper Maimaft
Gunnera macrophylla B1.,
HERNANDIACEAE
Hernandia ovigera L., SR, Pio; cf. CGS 758,
det. Croft
HY DRANGEACEAE
Dihvna ee.

WT 12183

Jebrifuga Lour., WT 12559, ‘sylvatica
complex’
ICACINACEAE
Citronella suaveolens (B\.) Howard, WT
11739, 36

Gomphandrar ‘a montana (Schellenb.) Sleamer,
AM 431; WT 11280, 11937
ae a Sp., ‘australiana-montana group’,
ogee pone peed ee srentiate
1.) Sleamer, WT 12652
var. borneensis (Heine)
&15, 1213

Platea oe Bi
tr, WT 1
aie ee (Pul le) Taine or

Papua New Guinea

971

dorae Moeser, WT 12771; flowers
red to differentiate
pene i ie ee ey
Wi ALGO; MOS, 222), 19,

L279, oe

Stemonurus monticolus (Schellenb.) Sleumer,
y O12
LAMIAC
Pisa alia (L.) R. Br., LAE
8, det.
EUs

nbelliflora (B1.) Hooker f., WT

i2 I 2
Betlschmtedea ct. schoddet Kosterm., WT
11198, 11814
C ees aff. aureosericea Kosterm., WT
1799
Perio cf. depressa Warb., WT 11922
Cryptocarya att. depressa Warb., WT 11862

Gr Lag aft, multipantculata Teschn., WT
202

Endioni ies Gamble. WT 13013
., WT 11920,

agen
1 2007
Endiandya ct. a nie i glauta
group, i ae eZ Sd Lae os
Litsea calophy nee - Hee WT 11699
ersea americana Mill., SR, Maimatu,
cultivated
LEEACEAE
Leea coryphantha Lauct., WT 11731
Leea (close to) heterodoxa K. Schum. & Laut.,
WT 12949; ‘coryphantha-heterodoxa group’
Leea heterodoxa K. Schum. & Laut., L
66334, Purari River; det. Leiden
Leeda ene rm. f.) Merr., re 486; WT
738, 12633
LEGUMINOSAE/CAESALPINIACEAI
se ii crista L., CGS 808, Purart, det.
_Le ‘iden

alate SR, Haia-Pio
: dia “pu ice , CES 804, Purari,

es: ee Kuntze, SR, Haia-
Pio; cf. S&C 4545, det. collectors
Kingiodendron sp., SR, Pio
ees schefferi ue Schum. & Hollrung,
2, Hata-Pio; cf. S&C 4406, det. Verdcourt

972

LEGUMINOSAE/FABACEAE
A ganohe ie (L.) Polhill, S&C 4464,
i, det. Verdcou
A ae es | Pee eee
Crotalaria lanata Beddome, WT 12018
G rOTATIESD Qoosbina ae Aiton, cf.
oe 8 - E. ste det. Verdcourt
rtl. CGS 830, Purari,
dee Verdcourt
Dalbergia beccarii Prain, S&C 4491, E. Purari,
det. Verdcourt
Derris cf. cunetfolia Grah. ex Benth. (sensu
Verdcourt), WT 11939
Derris ae Lour., CGS 815, E. Purati,

Desmodium a DC., WT 11773
Desmodium repandum Vahl) DC., WT 12244
12803
mesial ee es WT 12573, 12607
L.) DC., SR; cf. CGS
724, SEC pase a Leiden
Desmodium sp., (possibly) D. mM
L.) DC., cf. var. strigosum in CGS
S&C 4512
Gliricidia sepium Jacq.) Walp., SR, cul-
tivated

Inocarpus fagifer anaes Fosb., CGS 795,
iri, det. Leider
Inocarpus Sp. NOV.,
sensu Verdcourt,
Macropsychanthus lauterbachii Harms, WT
183°¢

abide morphospecies
T 11139, 12054

Mucuna aft. ?lamit Verdcourt, WT 11837

(fl), key uncertain eeu fruit
Muacuna novo-g SR; ct. LAE
6 Purart; SEC 4400

Pisa vulgaris L., SR, cultivated
“_e pinnata (L.) Pierre, CGS 828, 867

Purari, det. collectors
Pea He a now, SR; cf. SEC
C&S ,aeb collec
Strom er shia lecipie ner wrens
Were ee Hook. f., W7
Tephrosia sp., (possibly) vestita oe
cf. the sp. in CGS 905, det. ae

LEGUMINOSAE/MIMOSACEAE
Entada phaeseoloides (L.) Merr.,

logical

WT s.n.,

1
Mimosa pudica L., SR, Haia & Maimatu

Sipa 18(4)

Paraserianthes falcatarta (L.) Nielsen, SR,

cf. SGC 4242, det. Nielsen
ACEAE
mee mms Fv.M., WT s.n.
LOGAN

sae Sachin . Moore, WT 11695
Pagraed a teriana A. Gray ex Benth., WT
1279

Fagraed ae Thunb., AM 662
Fagraea elliptic SR, O-Pio
Fagraea ein we ee ex < Wall., SR, O-Pio
Fagraea woodiana Fv.M., SR, Wara Navarah
Gentostoma rupestre JR. & G. Forst., WT

907
era weinlandii K. Schum., WT
_ 12071, 1241
Neuba ve bode ee oe oe SEC
, Purari R., det

Nai noma oo AM
WT 1283

Nab 7 eas Leenh., WT
25,22 706

nae minor Bl., GW 799, det. GW; cf.
also SGC
LORANTHACEAE
\myema friesiana (K. Schum.) Danser, WT
12295
Amyema rigidiflora UX. Krause) Danser, WT
11869

Amyema squarvosa (IK. Krause) Danser, WT
12578, 12883
Dactyliophora verticillata (Scheffer) Tiegh.,

Decaisnina hollrungit US. Schum.) Barlow,
WT 12236, 12541

Dendrophthoe curvata (Bl.) Mig., SR, Haia

Macrosolen geminatus (Merr.) Danser, WT
12828

MAGNOLIACEAE

E pa cern (L. eae: ssp. ear
fampaca, AM 237, 392; WT 1

a
Magnolta candollii (Bl.) H. Keng var.
candollii, WT 12585, 12650
MALPIGHIACEAE
Ry a rys timoriensis (DC, var. discolor
sand.) Jacobs, WT 12564

TAKEUCHI, Plants from Crater Mc.,

MALVACEAE
Hibiscus archboldianus Borss., WT 11194,

12086
te L., SR; cf. CGS 730; SEC
522, det. olleérors
ae populnea (L.) Sol. ex Corr., WT
11971

Urena lobata L., WT 12459
MELASTOMATACEAE

Catanthera lysipetala Fv.M., WT 11302
Creochiton novoguineensts (Bakes f.) Veldk.

& Nayar, WT 12038
ie angiensis Ohwi, WT 11200,

ee sp., WT 11
— ies aff. albida oe & — WT

Pee aft. compacta Bakh. f., WT 11169
secre crassinervis Bl., AM 622; WT
, 12044, 13000

jee dentata Veldk., WT 12068

Medinilla ‘ramiflora group’, WT 12373; keys
t eld’s spp. 14-16

Medinilla rubrifructus Ohwi, AM 665; WT
11241

Medinilla sogerensis Bak. f., or aff., WT 12 |
eee ee Bak. Wi ii
Mail a smannit Mig., WT 12077,
ie aff. ?versteegi? Mansf., WT : 1114
Memecylon ‘hepaticum’ sensu ven Wr
12189, 12557; ke ee
Memecylon cf. ee eis Manst., W7
219, 11895, 124
Memecylon torricellense Manst., WT 12045
Ochthocharis bornensis Bl., SR, Pio; cf. S&C
4200, det. Hansen
Otanthera bracteata Korth., WT 12102
Otanthera cyanoides Triana, WT 11129,
12007, 12609
Mee multiflora Maxw., WT 12140,
2644 AA
Pile cf. robusta Mansf., WT 12034
2391-A, ‘robusta-macrophylla
BY i ne villosa Maxw., WT 11692

MELIACEAE
Aglaia (nearest) agglomerata Merr. & Perry,
AM 700

Papua New Guinea Pees)

Aglaia agglomerata Merr. & Perry, WT
11879, 12902, 12062
Aglaia argentea BI., WT 11956, 12025
Aglaia ee Hence WT 11851
Aglaia rimosa (Blanco) Merr., WT 12782
Aglaia sapindina pare oe WT 11187,
0
Aglaia subminutiflora C. DC., WT 11809,
11919
Aglaia cf. subminutiflora C. DC., WT 12882,
12956, atypical indumentum
Aglaia tomentosa Teijsm. & Binn., WT
11991, 12049
Aglaia aff. tomentosa Teijsm. & Binn., WT
11776

Aglaia sp. ?nov., aff. ‘grandis-ramotricha
9)

group, AM 6
Aglaia sp., (possibly) A. silvestris (Roemer)
Merr., cf. CGS 798, i

ere nitidula (Benth.) T.D. Penn.
T 11952, 12058
Aphananisis bolystachya (Wall.) R.N. Parker,

Chisocheton ee (Miq.) C. DC., WT
1916, 12060
Chisocheton lasiocarpus (Mig.) Valeton, WT
11966, 11998; entity ‘weinlandi?
a lastocarpus (Miq.) Valeton, WT
1938; entity ‘carol?’; = C. caroli Harms
A eu (C. DC.) Stevens, LAE
61 rari R., det. Stevens
ee sella Stevens WT 11975
1.) Mig SR: cf. CGS
788, Purari, det. Mabberley
Dysoxylum excelsum Bl., or af ‘alliaceum
rroup’, WT 171891, 11897, 12013
Dy ae gandichaudianum (A. de ) Mig.,
LAE 61125, Purari R., det Leiden
Dysoxylum tnopinatum ase s) acm WT

>

Dysoxylum cf. Raniense Harms, WT 11732,
2946
Dysoxylum mollissimum BL. ssp. molle (Miq.)
Mabb.,
Dysoxy bo papuanum (Merr. & Perry) Mabb.,

F lun par m(Osb.) Kosterm., WT
11733, 12174, "12954
a pettigrewianum FM. Bailey, WT
11923; LAE 6629

974

Dysoxylum ?randianum Merr. & Perry, or
aff., WT 12562
Dysoxylum setosum (Span.) Mig., WT
2591
Dysoxylum cf. spe elon WT 11829
Dysoxylum variabile Har W . 11694,
12021, 12291, 127 (be:
arms, WT 12/01
Vavaea amicorum Benth., WT 12103
Nylocarpus granatum Koen, , SGC 4484, E,
Purari, det. Leiden
sari ea moluccensis (Lam.) M. Roem., SGC

ari, as N. australasicus

12126,

Dysoxylum att. variabileH

MENISPERMACEAE

MONIMIACEAE
Kil Lbnlian

Arcangelisia tympanopoda (Laut. & K.Schum.)
iels, WT s.., carpological coll.
a aun ovate Mig., LAE 61110, Purari

wn

R., det. Forn
ei mninutiflore Cs. Schum.) Die
1948
Stephania dite hae: ) Miers var.
japonica, WT 12887
Stephania nomic (Thunb.) Miers var.
t/morte DC.) Forman, WT 120680
ae “ions Miq., WT 11212, 12464
Hook. f., WT 12096
Tinospora a poe & K. Schum.)
Di VT 11853

Tenancy ~p

iels,

A.C. Smith, LAE 66299,
Wabo dam site, det. Philipsc

<ibara ct. papuana A.C, Smith, WT
12728, ‘cortacea-papuana ae

[12722

Levieria a acuminata (F. v. M.)
Perkins, WT 12

Levieria montana a WT 12417

Levierta nitens Perkins, WT 12494

Palmeria brassti Philipson, WT 12595

Palmeria gracilis Perkins, AM 376
Palmeria gracilis Perkins var. ?nov., WT

12431; noncontormist indumentum

acres hirsuta (Warb.) Perkins, WT
12 ?] 4 29 2 3

Shae bnpiton ) Kanehira &
atusima,

Steganthera ue nov., val myrmecophilous
group’, WT 12

Stipa 18(4)

MORACEAE

Ficus adenosperma Mig., AM 209; GW 808,
det. GW

Ficus amblisyce Corner, AM 408, det. GW

Ficus ampelas Burm. f.,GW 713, det. GW

Ficus arbuscula beni & K. Schum., WT
2093

Ficus archboldiana Sammeth., GW 764, det.
GW

Ficus arfakensis King, GW 780, det. GW

Ficus augusta Corner, GW 791, det. GW

Ficus aurita Bl., GW 714, det. GW

Ficus bernaysii King, GW 712, 728, 784,

G

Ficus botryocarpa Mig., GW 753, det. GW

Ficus caesaroides King, Dodson s.n., det. GW

Ficus comitis King, pers. comm.

Ficus congesta Roxb., GW 720, 818, det.

GW

Ficus a Roxb. var. chalmersii (King)
forner, GW 7/1, det. GW

Freus es Steud., AM 501, det. GW

aha poner GW 805B, det. GW

iels, AM 273, 313, det.

GW; WT 5 5M.
Ficus dist Kae Diels, GW pers. comm.
Ficus edelfeltii King, Gw pers. comm.
Ficus erythrosperma NG. Ah det. GW
Ficus sal aian BL, WT]
Ficus gul Li & K. Mita oe 13014
Ficus habliane Diels, GW pers. comm.
: zeus homobroniana Corner, GW pers. comm.

1s homodroma Corner, WT 1182

a slid ue GW pers. comm.
TICHS IV GW pers. comm.
Ficus itoana Diels, wr 11121
Pi Laut. & K.Schum., GW

pers. comm,
Pic lophylla Diels, GW 776, det. GW
ers. comm,

WT 11679,

Ficus ene Lisi
Ficus mollior P.y.M. ex Benth.,
also GW pers. comm.
Ficus mollior ¥.v.M. ex Benth. v

Co , LAE 663 Purari River; det.

Var. SéS! salt. y

corner

Ficus nasuta Sammerh. var. glabrata Cor-
ner, GW 792, det. GW

Ficus nodosa Teysm. & Binn., WT 11987;
LAE 61098, Purari, det. Corner

TaKeucHI, Plants from Crater Mt., Papua New Guinea eye

Ficus ochrochlova Ridl., GW 735, 752, det. GW

Ficus pee King, GW 718, 782, 793,
det. / WT 11864

Ficus ovatacuta Corner, WT 11834, ?sp.
nov., does not

Ficus pachyrrhachis Laut. & K. Schum., GW
pers comm.

Ficus pachysycia Diels ex Corner, GW pers.
comm

Ficus pha Laut. & K. Schum., pers.

W based on JE 18 & 30 at A

Pet /la Diels a pers. comm,

Ficus polyantha Watrb., GW pers. comm

Ficus psendojaca Corner, a pers. comm.

— ee Reinw. ex BL, GW 717, 750,
det. GW

os ie Corner, GW

Ficus scratchleyana Kae, aioe. i G Ww

Ficus scratchleyana King var. rhopalosycia
(Diels) Corner, GW pers
on JE 27

Ficus semivestita Corner, GW pers. comm.

F 4 septica Burm. f., LAE 61

- comm. Dasec

125, Purari
., det. Cor
Ficus stervocarp Diels GW 709, 715, det. GW
Ficus sterrocarpa Diels var. pubigemma Diels,
GW

GW 754,

Ficus sterrocarpa Diels var. sterrocarpa, GW
5, det. GW
Ficus subcuneata Mig., AM =a sek GW
Ficus sublimbata Corner, Ww 3
Ficus subulata Bl., 779, av dee GW
Ficus subulata ne var. - gracillima (Diels)
er, GW 801, det. GW
Ficus tenella Corner, GW pers. comm.
Ficus ternatana Mig., GW pers. comm.
Ficus trachypison K. Schum., AM 536; GW
721, det. G
Ficus trichocerasa Diels, GW pers. comm.
Ficus virgata Reinw. ex Bl., GW pers. comm.
Ficus wassa Roxb., GW 796, det. GW
Ficus xylosycia Diels var. cylindrocarpa (Diels)
Corner, WT 11189

=

Ficus sp., sect. Conosycea, GW pers. comm.

Ficus sp., sect. Rhizocladus, GW pers. comm.

MYRISTICACEAE

Endocomia macrocoma (Miq.) de Wilde ee
prainii (King) de Wilde, WT 11958
12003, 12057

Gymnacranthera farquhariana (Hook. f. &

h.) Warb. var. zéppeliana (Miq.) R.
Schouten, WT 11908, 12050

ha ees: hellwigii (Warb.) Warb., WT

Hor: ie i (Bl.) Warb., ee ! sree
Horsfieldia pulverulenta Warb., 195
Horsfieldia subtilis (Miq.) aa var. ia
2 WT 11883
Horsfieldia subtilis (Miq.) Warb. var. aucta
de Wilde, WT 11674
Horsfieldia sylvestris (Houtt.) Warb., WT
s.n., sterile
Hor aa . oe CK. Schum.) Watb.,
paaee possibly H. hellwigii
aJ.Sinclair, WT 11261,

Myristica cf. cucullata basa id J oo
Myristica globosa Miq.,
Myristica aff. globosa Miq., ee ae
Myristica ingens (Foreman) W.J. de Wilde,
WT 10251
Myristica cf. inutilis Rich. ex A. Gray subsp.
papuana (Markgraf) W.J. de Wilde, WT
11865,
Myristica ube Mig., WT 11262

MYRSINACEA

Ardisia aft. squarrosa nae WT 11281,
11904, 12076, det. Pipe

Loberia veiniana bs See WT 11921

Maesa haplobotrys Fv.M., WT 11855

Maesa ee Fv.M. ae a 30, Purari
Ri

Wes aff. acrostica (Mez) Pipoly, WT

an pee sp. nov., det. Pipoly
/ (1S. Schum.) Pipoly, WT
| 1693, det. Pipoly

MYRTACEAE

Acmena acuminatissima (Bl.) Merr. & Perry,
/T 12100
Decaspermum bracteatum (Roxb.) A.J. Scott,
R, Hata & Pio; cf. S&C 4373
Seis mum exiguum Merr. & Perry, WT
< 58 <<
Decprma ee tlum Laut. & K,
Schum., AM 625; WT 11216
Merida iit Laut., SR, Maimatu,
alsc t. by Maimafu miles

976

Octamyrtus bebrmannit Diels, WT 11677

Octamyrtis ees os M.) Diels var.

plewopetata, Wl 4 (fr), 12617 (ft),
12759

sii latifolia (Benth.) Mig., WT
3]

a novogiineensis Diels, WT 12421

ulum K. Schum., WT 12076

Syzygium buettnertanum (&. Schum.)
Niedenzu, SR, Maimatu, possibly S

Syzygium acutangu

subcorymbosum
Syzygium callianthim Merr. & Perry, WT
12193

S yey etm decipiens (Noorders & Valeton) Merr.
tebe ee oi T 11767, 12118
Syyei 1 Merr. & Perry, WT

}

112 Dar arf
Syzyerum effusum (A, Gray) C. Muell., WT
12173, 12844
Syzygium Geen Merr. & Perry, SR,
Maimafu
sie Cre eee & Perry,
VT 12525, ‘hylochare-phaeostictum ee

Syzyeium plan Diels, or aff.,
12698,

SyZ zy giteme ons Merr. & Perry, WT 12655
Syz3 (L.) Merr. & Perry, sensu
lao, WT l sie. i ty 7, 12940, 12944
Syzyeu vault. & K.Schum.)

Mee & soi Wr 11676, or aff.
megalospermum
Syzyeuem porphy ea es ves) Merr. &
Perry, WT 12000, 12497-A, 12877
Syzygiune aff. rosewm Mert, & Perry, WT
11847, po

ssible sp.
Sy?) 2 glum oe ahi. ge tatu (ih ) Merr.
rry, or = WT 11 11719

1 a4; ?ne

Syzygium Ae (Ridl.) Merr. & Perry,

Syz Seal ‘trerneyanum group’, SR, Haia&
Maimafu, with diffuse inflorescence

Syzygium trachyanthum (Diels) Merr. & Perry,
or aff., WT 11990

Syzygium tympananthum (Diels) Merr. &
Perry, WT 12598

Syzygium aff. womersleyi eee! & Perry,
WT 112

Nanthomyrtus spp., SR, Werner: fu, sterile,

258, possible sp. nov

more than one taxon

Sipa 18(4)
si ~ r sie AE
i Nees, WT 12284, 12319
A lava cf. papuana Dans., WT 11779

NYCTAGINACEAE
Pisonia ue Teysm. & Binn., WT
1201
— onli (Forst.) Seem., LAE
, Purari, det. Hyland
OCHNACEAE
Schuurmansia henningsii K. Schum., WT
12129
OLACACEAE
Anacolosa papuana Schellenb., SR, near Pio
Olax imbricata Roxb., CGS 829, Purari
OLEACEAE
Chionanthus riparius (Lingelsh.) Kiew, WT
11909

Chionanthus ?sp. nov. at
(Lingelsh.) Kiew, WT

C oe sesstliflorum ( ferisl ) Kiew, WT
292

salicifolius
905

¢ ssh an atf. Lifl (H lems sl.) Kiew,
WT TLL75
uae sp. nov., WT 11866
ONAGRACEAE
Ludwigia . ssopifolia (Don.) Exell, LAE

96357, det. Kerenga
Ludwigia octovalvis (Jacq.) Raven, WT
1175 Z
PASSIFLORACEAE
a foetida L., a - ala
mafu; cf. SGC 4
PENTAPHRAGMATACEAE
Pentaphragma grandiflorum Kurz, WT 11163

1 & lower

PIPERACEA
Peperomia eS eee: ae a aaa
group, W711 nota
Peperontia ae ss .) Mig., WT 12342
a omia pellucida (L. is Kunth, SR, Haia
Piper me vm L., SR,
oo SR, cated
Piper caninum Bl,
a ail O; IZ, ba a 7 12100,

212
eibbilinbum C. DC., WT 12393,
55; hirtellous form
Piper gibbilimbum C. DC. WT 12268; typical

glabrous form

TAKEUCHI, Plants from Crater Mt.,

Piper macropiper Pennant, WT 11683, 12109
Piper majusculum Bl., SR, Pio; cf. SGC 4331
. Leiden
Piper mestonti ee WT 12879;
Stenocarpun
Piper novo-guineensis Warb.,
ak ae K. Schum. & Laut

‘form

WT 12090
WT 11681,

Piper laa K. Schum. & Laut., AM
5 1143

Bite, ie ei C. DC., WT 11741
Piper triangulare Chew, WT 12211, 12273,
12553, 12892
ge umbellatum L. var. fans
Willdenow) C. DC., AM 655; W
1224
Piper versteegit C. DC., WT roa
Piper w pees Chew, SR

Piper wv., WT 12453; ne filiform
ene
PITTOSPORACEAE
Pittosporum ferruginenm a f. subsp.
ferrugineum, WT 1267

ae pullifolinm am WT 11285,
74
49

Se pullifolinm ssp. ledermannii
ritzel) Schodde var. ledermannit, WT
12539, 12580
Pittosporum sinuatum Bl., WT 11153, 11279
Pittosporum stnuatum Bl. var. sinuatum, WT
POLYGAL
Ents ee papuana J.J. Sm.,
Maimafu
oi ap ne L., WT 12265-B
Securidaca ecristata Kassau, SR, Haia
a eat ad papuanum Whitmore ex
Meijden, WT 11764, 12032
POLYGONAC
oo, a L., WT 12009
PROTEA
Helicia of ee C.T. White, WT 12035
RANUNCULACEAE
Clematis papuasica noes & Ries SR, Pio;
> 4399, det. Tamur
Clee: phaner aii vere & Perry var.
simplicifolia Tamura, WT 12390, 12567,
12780

Papua New Guinea

S77

Clematis he Merr. & Perry cf.
var. tomentosa Eichler, WT 11256
eee sp., SR, upper Maimafu
RHAMNAC
Apion ine (Fenzl) Reiss. ex Endl.,
T 12218; sensu Schiraren
Alphitn smacrocarpa ene i. Pio
., SR, Haia
Cue javanica Miq., es oe
Rhamnus ping (Wallich) ain ex
Hook., WT 11690, 12,277, 12674

1271 6 ; 27d -
Ventilago ?papuana Merr. & Perry, SR, Pio
Zizyphus angustifolius (Miq.) Hatusima, WT
11985
Zizyphus djamunensis Laut., LAE 66318 near
o, det. Frodin

RHIZOPHORACEAE
Carallia a (Lour.) Merr., SR, Pio;
C

Gynotroch wis Bl., AM 428, 522; WT
12089
ROSAC

EAE

Prunus arborea (Bl.) Kalkman, SR, Haia

Prunus dolichobotrys (Laut. & K. Schum.)
Kalk aia

Prunus gazelecpeninsala (Kanehira &
Hatusima) Kalkman, WT 12642, 12661,
1 2858

Prunus oligantha Kalkman, WT 12802

Prunus schlechteri (Koehne) Kalkman, AM
328

Rubus moluccanus L. var. moluccanus, WT
12022
Rubus rosifolius JE Smith, WT 12095

RUBIACEAE
Airosperma aff. ramuense Laut. & K.Schum.,
WT 115

sae brassii Merr. & Perry, WT
2908

Amara aff. grandifolius Valeton, WT
12495

Antirhea sp., Se eae Merr. &
Perry, OlCE SI t. Darwin

>

Argostemma ee K on um., VAM Dies
WT 11132, 11150

Argostenma callitrichum Valeton, WT 12364,
12?

978

Canthinin aft. cymigerwm (Valeton) B.L. Burtt,
VT 11868, 12128, 12737, not the

1es
Canin cf. longifolinm (Valeton) Merr. &
7T 12106-R
pre sp., WT 12934; also diff. sp.
in SGC 4233, Gult
Dolicholobium gertrudis K. Schum., WT
12110, 12498
Dolicholobinm oxylobum '&. Schum., WT
11965

Gardenia hansemannit &.Schum., SR, Pio;
ft. CGS 768

Gardenia lamingrni M. Bailey, SR, Haia;
388, det. Leiden
Ses Merr. & Perry, WT 12399,

12447, 12479, 12829
Hedyotis auricularia L., WT 12666
Hedyotis schlechteri (Valeton) Merr. & Perry,
YT 11820

ae radicans Becc., LAE 61087,
iR., eiden
Ip terrestris Huxley & Jebb, or
"12587, 12831
ae on atts or aff., WT 11926
xora aft. moszkowskis Brem., WT 11936
Lastanthus chlor carpus K. Schum., or aff.,
WT 12410, 12734
Lasianthus tomentosis Bl., WT 12158, 12220,
2732
a (closest to) fomentosus BL., WT
12885, 12894
ee ee ied K.Schum. & Laut.,
or atf., WT 11213; = M. simplex Merr.
& Perr
se rag On ee (kK. Schum.)
SGC 4278, Gulf D., det.
Darwin
Morinda citrifolia L., CGS 821, E. Purari,
et. Gideon
Morinda umbellata L. var. papuana Valeton,
WT 12058

Mussaenda bevant ¥v.M., AM 214, 379; GW

723, det. Gideon

Mussaenda ferruginea &. Schum., WT 12049

Mussaenda oreadum Wernh., WT 12615,
det. Gideon

Mussaenda scratchleyi Wernh., SR; cf. SEC
4238, 43006, det. Gideon

ae]

Stipa 18(4)
Mycetia javanica (Bl.) Reinw. ex Korth.,
82, 11221
Myrmecodia platytyrea Becc. subsp. platytyrea,
WT 11193

Myrmecodia sp., SR, Maimafu near clinic,
not platytyrea

Nauclea orientalis L., GW 803, det. GW

Neonauclea acuminata Ridsd., WT 12055

Neonauclea gordoniana (F.M. Bailey) Ridsd.,
WT 124

Neonauclea cf. perspicuinervia Merr. & Perry,

Ophiorrbiza att. amoena Valeton, WT 11271,
12088
Ophiorrhiza atf. ?debrnynit Valeton, WT
12010, 12084, 12246-B
Ophionbiz elif Val eton, i 11149
Ophiorrhiza sp., unplaced af [12411
Saale guele ae K. eice. AM
541; WT 12160, 12721

Psychotria amplithyrs
12900

Psychotria att. amplithyrsa Valeton, WT
12028, vining, not the sp., not in Valeton

ie ct. chrys antha Merr. & Perry,

er ae 12070, 42102,

/ 2891, gon to fork: 44

Psychotria dieniensis Merr. & Perry, WT 12343

Psychotria ct. hollandiae Valeton, WT 12014

Psychotria leonardii Mert. & Perry, or aff.,
WT

Valeton, WT 12099,

12960
a ihe aff. /eptothyrsa Mig., WT 12662
ton key, not the s
Pssebotria mucralabastra (Laut. & K. Schum.)
Valecon, LAE 6/167, Purari River, det.
ohmer

Psychotria multicostata Valeton, WT 11152

Psychotria myrmecophila Laut. & K.Schum.,
AE 66280, Wabo dam, det. Sohmer
WT 11765; cf.

Nilaw!
UY

~

Psychotria oe ta i iuleton,
7, Gulf
Psychotria aff. ae Valeton, WT 12397
Psychotria purariensis Sohmer, LAE 66298,
66349, det. Sohmer

P : Las la ramadec umbens So
. 12904

amer, WT

oe cf. ramulosa Merr. & Perry,
‘micrococca group’, WT 12035, keys to
77-78, stipule ramulosa

TAKEUCHI, Plants from Crater Mt., Papua New Guinea D9.

Psychotria sphaerothyrsa Valeton, WT 11678. _ Sia Re seerel laevicar di end
12979 SR, Hai

Psychotria valetontana Sohmer, WT 12817, eo dy ee R. v. ove oe WT 11857

2843, 12893 Lunasia amara Blanco var. amara, WT s.n.

Psychotria womersleyi Sohmer, WT 12028, Micromelum minutum (Forse. f.) W. & A.,
12056 SRect, SGC 4542) CES 1003, Gulf

Psychotria sp. nov., aff. eee Takeuchi, Tetractomia i (Roxb.) Merr., WT
WT 12130, 12602-B, 12285, 12594

Psychotria sp., unplaced e cee 12706 Venzelia tenuifolia Swingle, WT 12958

‘Randia’ decora Valeton, W T 11915, ge- pin oaiiecaiste ten i Merr. & Perry,
nus name is problematic WT 12465

‘Randia’ schumanniana Merr. & Perry, WT nee lum spp., SR, Hata-Pio

22 aaa pa Roxb )DC., SGC 4246,
‘Randia’ sessilis Ev.M., WT 11917 "Cal

Rhadinopus papuana S. Moore, WT 11630 (possibl a ae: Wighe, C&S 1019, Gulf
‘Tavenna buruensts (Miq.) Valeton, WT 11684 SABIACEAE

Timonius belensis Merr. & Perry, AM 415; vf ta Roxb.) Maxim. ssp. humilis

Merr. ‘& Perry) Beus., SR, upper Maimafu
Meliosma pinnata (ob. ) Maxim. ssp.
ba fla (Merr.) Beus., WT 11190,

Timonius grandifolius Valeton, WT 11925,

Timonius timon (Spreng.) Merr., GW 758

Timontius trichanthus Merr. & Perry, WT Pr ve iflora Bl., WT 12226, 12959,
2187, 12548, 12808
Timonius sp., ‘laevigatus-pulposus group’, WT
< SANTALACEAE
ae : ; xa aft. unci anser, WT 1235¢
Uncaria lanosa Wall. var. oo Cladumyza ais wamanata Vonset WI 12522;

(Benth.) Ridsd., AM 378; WT 11199 keys to wncinata but not the sp.
rieaeis wens Rie WE 12008 Dendromyza reinwardtiana (B\.) Danser, WT
: * 12348

Urophyllum aff. 2brittanicum Wernh., WT

Dendrotr ph pl: St auffer, WT 1 1856
Versteegia minor Valeton, WT 11188, me Sana Jivanteen: (Laur. & K.Schum.)
7 9
iss sas es )DC., Pilg., AM 679; WT 11253, 11707
SZ, 1252 SAPINDACEAE

Page / ed monospermous shrub Alectryon myrmecopbilus Leenh., WT 12746
RUTACEAE Allophylus cobbe (L.) Racicchel SR, Pio;
Acronychia sp., SR, Haia cf. ne sie det. oe ese
Exodia hortensis J.R. & G. Forst., WT 12490, Cup scropetala Rac 11146

Cian aie Radlk., ID based on
ruits on ground, Haia
12280, 12636 Cupaniopsis ee oe as : 12864
Melicope grandifolia B.L. Burtt, or aff., WT Dictyoneura obtuse
Dodonaea angusti ae if - ae
a uioa molliuscula Radlk., or a WT ae
299)

Melicope denhamti (Seem.) T. Hartley, or aff.,
: 26

8

Lio¢1
Melicope (probably) mucronata Merr. & Perry,

R, upper Maimafu a, } Radlk., WT 12276, 12283
Melicope cf. nova-guineensis Valeton, WT MATTE SA - pair

fo ey 16 here bean econhem ow arborea (Blanco) Radlk., SR; cf.

Melicope sy , as Exodia §§ Coriaceae, SR; 4641, Gulf
cf. SEC S1¢ ree camptonenra Radlk., WT 11889
Flindersia ee Poir., SR; cf. SGC Harpullia ct. camptonenra Radlk., WT 12723,

4656. Cull det, Leiden possibly H. cupanioides

980

Harpullia longipetala Leenh., LAE 61175,
rari R., dec. eciden
i sehunltia pork

pet

ee (close to) subsp.
nee WT 12922

Harpullia sp., (prob. on ) ramiflora Radlk.,

Taia-Pio
es ee (BL.) Kalkman ssp. javanica,

be ween Radlk., WT 11950,
988, 11989
Lepisanthes senegalensis Leenh., SR, Pio; cf.
S&C 4337, det. Leiden
Mischocarpus ie eige cae , AM 597
Maschocarpus pyriformis (E. Radlk. subsp
retusis (Radlk.) R. a one WT 12 249,
2443

Maschocarpus sp., WT 12589
Pometia pinnata Forst., WT s,
Sarcotechta crispata Welsen. WT 11885
genus indet., WT 12413, 12694, 12784;
alate rachis
SAPOTACEAE
Pouteria anteridifera (C.T. White & Francis)
Baehni, AM 221
SAXIFRAGACEAE
Quintinia sp., SR, upper Maimafu
SOLANACEAE
Capsicum annuum L., SR, cultivated Haia
& Melman
Lycof lentim Mill.
a as L, sire SICH
Nicotiana tabacum L.., SR, cultivated
Physalis peruviana L., WT 12301
WT 12015
Solanum anfractum Symon, WT 12912,

28

, SR, cultivated,

Solanum americanum Mi ill.,

Solanum biflorum Lour., WT 12508, | 20
Solanum cladotr ae Bitter, WT 12
Solanum cf. nase esas Warb., wr
12705; smoalabastrum
Solanum dendroptlosum ee WT 11854,

5 possi

t. Symon

SR, Pio; cf. SEC

m Gaud.,
4244, det. Symon
Solanum bee vanum 1K. Schum. & Laut.,
CGS 848, Purari, dec. Symon
I,

Solanum per m Wernh., WT 11204,

cont. aon

Stipa 18(4)

Solanum pustulatum Symon, WT 11704,
onf. Symon

Solanum ct. pustulatum Symon, or aff., WT
12085; does not ae out

Solanum torvoideum Merr. & Perry, WT 12549

Solanum sp., (possibly) es Merr. &
Perry, cf. CGS 714; S&C 4625

SPHENOSTEMONACEAE

dets. Symon

Sa ce paloe Sane ) Steen. &
rdtman, WT 12444
S Paha ACEAE
andra (Schler.) v.d. Linden, AM

Wage WT 11858, 12119, 12432, 12478
STERCULIACEAE
Brachychiton carruthersii F.v.M., or B.
velutinosus Kosterm., WT 12021
Commersonta a (L. oan SR, Hata
Melochia odorata L.£., WT s.n., partial a
Pre saci me 17K, a ,CGS 77
WT 5.n., cé pion tee coll.
r.) Kosterm., SR, Hata
oe ps Bak. : ,WT 11186, 12056
Sterculia gilva Miq., or aff., WT 12814
Sterculia macrophylla Vane. WT 11984
Sterculia monticola Mildbr. var laxiflora Tantra,
T 7

Pr }

< O, a

— schumanniana (Laut.) Mildbr., SR,

a _ awit é v.M. var. shillinelawii,
WT s.n., partial s
Sterculia eet a: ae S&C 4467, E.
U ., det. Tantra
STYRACACEAE
Bryzycnyiey chyy,

fes Boerl. & Koorders, WT

Styrax arereste (Lour.) G. Don, SR, Pio &
ower Maimatu
SYMPLOCACEAE
Symplocos cochinchinensis (Lour.) 8. Moore
Leptophylta (Brand) Nooteb., AMI 659;
W

2

1233

‘THEACEAE

Adinandra forbesii Bak. f., WT 12120
Eurya att. kanehirai obusid: WT 11848,
-o here but leaf not crenate

— tigang WK. Schum. & Laut., WT 12063,
299

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 981

Gordonia papuana Kobuski, WT 12767 ee beccarit Schroter, WT 12031,
Ternstroemta britteniana F.v.M., WT 12405 2152
‘Ternstroemia wae (EM. Bail.) Merr., WT’ 5.7. Peas! blechnoides Ridl., or aff., WT
THY MELAEACEA 121 153 =F r Mae tule H. Winkler
Phateria epee (Scheffer) Boerl., SR, Elatostema cf. macrophyllum Brongn., WT
Haia; cf. LAE 61132, Purari R 11229, 12090, 12246-A
Phateria pilistyla Stevens, WT 12 069 Elatostema att. macropbyllum Brongn., WT
Phaleria sogerensis 8. Moore, WT 11808, 12584
11888, 12182, 12659 Elatostema mongiense Laut., WT see
Phateria cf. sogerensis S. Moore, WT 12786 Elatostema morobense Perry, WT 12

Elatostema ct. novognineense Warb., W r i we 9

Elatostema polioneurum Hall. f. SR Maimafu

Elatostema tridens Perry, WT 12385

i decumana (Roxb.) Wedd., WT
936

TILIACEAE
Grewia aft. brassit Burrett, WT 13017
Maicrocos ct. grandiflora Burrett, WT 11903,
12004

Aicvacos tetrasher LE 7)
ih ioe alles ee pe hetty, WT LLOS2 Lali oon ) Wedd., SR, Pio;
Trichospermum tripyxts (XK. Schum.) Kosterm., pelaeed pecs

y 2? ? ; ie y '
WT 12366, 12743 Nothocnide on ep Chew, WT 12389

Nothocnide repanda (B\.) BL., S&C 4501; WT
746

Triumfetta sp., ‘nigricans-ptilosa group’, WT
12396

TRIMENIACEAE Oreocnide trinervis (Bl.) Mig., AM 527, 618;
Trimenia papuana Ridl., WT 12857 WT 11177. 12681

ULMACEAE Pilea caudata Winkler, WT 12254
Celtis hildebrandii Soepadmo, WT 12095 Pilea cuneata Winkler, sensu Chew, WT

12279
Pilea effusa Winkler, WT 12175, 12529
Gironniera (probably) Airta Ridl., SR, a Pilea papuana Winkler, WT 12267

Celtis sp., ‘latifolia-philippensis group’, WT
s.m., sterile

Parasponia rigida Merr. & Perry, Ww’ LT 12206 Pilea venulosa BI., WT 12117

Parasponia rugosa Bl., WT 12080, 12 260 Pipturus argenteus (Forse. £.) Wedd., WT
Trema cannabina one WT 11838 11806, 12520, 12627

Trema orientalis (L.) Bl., SR, Haia Pouzolzia sp., SR, Maimafu

URTICACEA
Cypholophus aft. decipiens Winkler, WT 12384
Cy ae as latifolius (Bl.) Wedd., WT

— es (Winkler) Schréter, WT
12

Procris ai (Forst.) Wedd., WT
2039
ALG ol. . 7
Clb cf. pachycarpus Winkler, WT Pr 2 gE Ee
Copbolopbes rotundifolius Winkler, or aff., ©. VERBENAC
caliane onsifolia Lamk., WT 12412,

c ae cf. velutinus Winkler, WT 12823;

kevs to velutinus on Winkler calla Dees R. Br., WT 12003
Daigs longifolia (Burm. f.) Wedd., WT 77 Beer & HJ. Lam, AM
217; WT 11227
De ee sp., WT 12207, 12846, 12868 Clerodendrum buruanum Mig., WT 11297
eae a (BI.) Chew, LAE 61082, Clerodendrum inerme (L.) Gaertn., SR, Pio;
S&C 4474
Late sae ternatensis (Miq.) Chew, WT Clerodendrum ?sp. nov., WT 12209; no result

12019 on Lam, no LAE match

982

— splendida F.v.M., SR, Pio; cf. CES
urari
a eae (Roxb.) Merr., WT 12425
Gmelina dalrympleana (P.v.M.) H.J. Lam,
SR, Pio
Premna serratifolia L.., SR; cf. C6S 818, E.
Purari
Teijsmeanniodendron abernianum (Merr.) Bakh.,
JT sn.
ar cofassus Reinw. ex BL., SR; cf. SGC
497, Purari

VITACEAE

Ampelocissus sp., SR, Haia
Cayratia japonica (Thunb.) Gagn., WT
Il

Cayratia (close to) japonica (Thunb.) Gagn.,
WT 12108

cf. CGS 855, Purari, indec.,
‘trifolta facies’

Cissus avistata Bl., WT 11759

Cissus sp., aff. aristata Bl., WT 12094,

2985 t the speci

Cissus ieee ea or cout SR, Haia;
cf. S&C 4505

Nothocissus penninervis (F.v.M.) Lacciff, LAE
61100, Purari R., dec. Leiden

Tetrastioma sp., WT 12456, 12645

Cayratia sp.,

WINTERACEAE

Zyvogynum aft. 2elaucum (A.C. Smith) Vink,

268

MONOCOTS

AGAVACE

EAE
a ct. fruticosa (L.) A. Chev., WT

AMARYLLIDACEA

Crinum asiaticum a LAE 61090, Purari
R., det. Leiden list
Curculigo capitilata (Lour.) Kuntze, WT

1088
Curculigo cf. ciaied (Lour.) Kuntze, WT
2789; inflorescence
erect

ARACEAE

Ani bial en SR, Pio
L.)G. Don, WT 11296
Alocasta nicolsonit A. Hay, WT 12202

SIDA 18(4)
alae cf. nicolsonii A. Hay, WT 12145,
12859
Amupophals pe EM. Bailey, WT
Petes ae (Schott) Nicolson,
WT 12048

Colocasia esculenta (L.) Schott, SR, cultivated

G rane ciliata a ) schote, Sac

. Purari R., det. N. Jacobsen

cn et. a Alderw., WT
208

ce pinnatum (1..) Engl.,

Holochlamys beccarii Engl.,

Pothos falcifolins Engl. & K. Krause, AM
509, det. A. Hay

AM 600
)

ARECACEAE et ae

Areca catechu L., SR, cultivated
Calamus hile Bece., WT sn.
Caryota dose Martelli, WT 12665
Cocos nucifera L., SR, cultivated

Metroxylon sagu are. SR, Haia

COMMELINACEA

Aneilema cf. acuminatum R. Br., LAE 66362,
Wabo

ae diffusa Burm. f., SR,
Floscopa scandens Lour., LAE ae ‘Wabo
esse mollissima (Bl.) Koords., s. lat.,
[ 11863, = ?Amischotolype
os ere (BI.) Steud., WT 11784

COSTACEAE

Costus specrosus (Koen.) J.E. Sm., SR, Hata
ce bo ANANASSAE ci ) K. Sc hum.,
855 ideon
ea brcisti Gideon, pers. comm,
sideon
ala versteegii Valeton, AM 596;
VT 11156, 11179, det. Gideon

CY PERACEAI

Cyperus cf. nutans Vahl var. efewsrnordes

(Kunth) Haines, LAE 66361, Wabo, det.

Leiden

aramapania parvibractea (Clarke) Uittien,
LAE 66307, Wabo d site, det. Henty

Suede mess eae WT 1174 7

, WT 12686, 12787

DIOSCOREACEA

Dioscorea bulbifera L., WT 13006

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 983

Dioscor ea (probably) escw/enta (Lour.) Burk.,
. io & lower Maimafu
Deine cf. nummularia Lamk., WT 11768,

FLAGELLARIAC
Flagellaria eae . WT 12216

GEITONOPLESIACEAE
Geitonoplesiam oe Br.) Hook., WT

HELICONIACEAE
Heliconia papuana WJ. Kress, WT 11882
IRIDACEAE
Tritonia x crocosmiflora (Lemoine) Nichols,
WT 12807

LILIACEAE
Dianella ensifolia (L.) DC., WT 12287,
82

Dianella javanica (B\.) Kunth, or aff., WT

MUSACEAE
Ensete aan Lies ) Cheesman, SR,
yerb, Maimafu
ae wae a WT s.n.

RCHIDACEAE (dets. N.H.S. Howcroft)

Bulbophyllum sas Schltr., He 38
Bulbophyllum cf. cavibulbum J.J. oa ,He8
Bulbophyllum intersiti ee erm acl: Hel
Bulbophyllum lepanthiflorum Schltr., Ha 1
Bulbophy oe (close to) nematapodium, He
58)

Bz lhophylli m pt dacanthy, my.J. Vermeulen,

He 37
Bulbophyllum trifilum subsp. trifilum J.J.Sm.,
WT 12810, 12903, 12975
Bulbophyllum wakoi N.H.S. Howcroft, type
COLE.
Calanthe ventrilabrum Rchb. f., He 21
Calanthe werneri Schltr., er ee ai oi)
Coelogyne beccarit Rchb. f., WT 12774
Coelogyne fragrans Schler., WT 12536, 12979)
12824

Coelogyne cf. fragrans Schler., He 13, 30;
Ha 8

oo veratrifolia (Reinw.) Bl., WT
240

Cystore - cf. dentifera Schltr., WT 11165
De bulbophylliodes Schltr., WT 12825

we faa cf. poole i as
WT 12872 1ucho

Dendr en nie Schier. Ww 12967

oe obinm ees Schltr., WT 12812
ndrobinm Cf. Sea J. Sm., §
valyptro chi ilus, WT 12290, 12372

Dendrobium pleianthum Schltr., He 33

ees wn se ase ual Reeve and J.

eae eee J.J. Sm., WT

Dendrobium rupestre J.J. Sm., He 17

plan aff. esis Rolfe, §

yptrochilus, WT |

Dends ies to En Krzl., §
Oxystophyllum, WT /2357

Dendr obium aff. vexillarius J.J. Sm.,

Oxyglossum, WT 12598

Dendrobium sp. A, § Calyptrochilus, He

16, 28

Dendrobium sp. — S Grastidium (mixed
l.), WT 7

Dandi ES c - Gram, WT 12986
BE oe Ze WT
on 12229, ae 12988
es chrysotropis Schitr., WT
3), 12298

ie sp., 856
Dipodium pandanum ai oe Foe LAE
66276, Wabo dam site
Epiblastus sp., WT 12045
Eria imbricata J.J. Sm., He 36
Eucosta papuana Sci alec WT 12136
Glomera sp., WT 12376
Paice or Peristylus, sp. A, WT 12101
Habenaria ot Peristylis, sp. B, WT 12195
Habenaria or Peristylus, sp. C, WT 12484
Habenaria or Peristylus, sp. D, WT 12586
iparis caespitosa Lindl., WT 12327
ae aff. gibbosa Finet, He 2
Distichum, WT 12

’

Liparts sp., §
Mail quadrifolium Schuipman, WT

es sp. A, WT 12446
Mediocalcar sp. B, WT 12815

Nenwiedia veratrifolia Bl., WT oe
Octarrhena angraecoides Schltr

Peristylus dolichocauton (Schltr. ) ee WT

984

Phaius montanus Schltr., WT 12

Phaius ct. montanus Schlte., WT
12826

Phaius sp., WT 12286

Phreatia ?canlescens Ames, WT 12297

Phreatia elata Schitr., WT 12933

Phreatia sp. A, WT 12794

Phreatia sp. B, WT : le

Oberonia sp., WT 12

Robiquetia cf. mooreana eciege: Sm., WT

7 2760,

Spathoglottis panlinae ¥.v.M., WT 12289

Spathoglottis plicata Krzl., WT 11286

Spathoglottis plicata . ed puberila N.S.
Howcroft, WT

Vanilla wariensis sh . T 11842
Vrydagzynea cf. all ler, WT 12385
Vrydagzynea cf. albostriata Sek ler., or
salomonensts Schltr., uty 12251]
Vrydagzynia sp.. WT 12341

PANDANACEAE

Frey, anetld LHL SEASS IML 1 Ridl.,

SR of com-

mon climber
Freycinetia att, 2lagenicarpa Warb., WT 11761
Freycinetia mavantifolia Hemsl., WT 11955
Pandanus leptocarpus Martelli, S&C 4493,
Purari, cited by Stone (1987)

Pandanus spp., WT s.n.

POACEAE (GRAMINEAE)

Apluda mutica . a cf. CGS 1065

Centotheca ena cee ) Trin., WT 11717

Chrysopogon acicitlatus (Retz.) Trin., SR; ef.
SEC 4593, det. collectors

Cotx lachryma-jobi L., WT 12019

Cyrtococcum accrescens (Trin.) Stapf, WT 12639

Garnotia stricta Brongn. var. longiseta Hack.,
WT 12470

Ichnanthus vicinus (EM. Bailey) Merr., WT
12717

SIDA 18(4)

shia inane WT 12408
LJP , WT 12599

12054,

Is ne iby
12059, 2064
aaa myosotis Nees, WT 12505
1s longispicula Holttum, WT 12600
oni bed (L.) P. Beauy., WT 12002,
715
Benne sp
Brongn.) at ri
Saccharum edule oe ie = ee
Saccharum officinarum L., SR, cultivated
Saccharum sp., ‘robustum - spontaneum com-
slex’, WT 12597
Scrotochloa urceolata (Roxb.) Judziewicz, SR
Pieaies wee Giger ) Stapf, WT 12006
Zea mays L., SR, cultivated
TRIURIDACE
ere lin Schltr., LAE 66347, det.
Leiden lis
Sctaphila es BL., LAE 66268, Wabo dam
site, det. Leiden

na WT

eae matcrostal hyum
5 natu

ZINGIBERACEAE
Alpinia werneri Valeton, § Pycnanthus, WT
1787
Alpinia sp., § Dieramalpinia, WT 11831
Curcuma australasica Hook. f., AM 382;
WT 11295

Etlingera dekockii, or E. versteegit, WT 11705;
d flower color
Hornstedtia scotttana (F.v.M.) IK. Schum.,
VT s

Phryniuin aff pedunculatum Warburg, WT
1880
ae aol macropye cilia (Valeton)

nich, WT 1

via
Pe ee ie. se 11141
Riedelia macrantha K.Scham., WT 11173

ACKNOWLEDGMENTS

The botanical survey of Crater Mt. was made possible by principal fund-
ing from the Liz Claiborne and Art Ortenberg Foundation. Financial sup-
port for the field component was also provided by the John D. and Catherine
T. MacArthur Foundation. Staff of the Research and Conservation Founda-
tion of PNG (Robert Bino, John Ericho, Paul Hukahu, Paul Igag, and Arlyne

—

Johnson) assisted with

1

ogistics and community liaison.

TAKEUCHI, Plants from Crater Mt., Papua New Guinea 985

The Lae National Herbarium served as the principal logistical base and
provided facilities for processing and identification of collections. N.H.S.
Howcroft illustrated the new species and colleague J. Pipoly of the Botani-
cal Research Institute of Texas corrected the Latin diagnoses. Keiko Muto
provided the Japanese translation.

A number of taxonomic specialists made plant identifications or assisted
indirectly by generously sharing unpublished information on their respec-
tive groups. My thanks are extended to B.L. Burtt (Gesneriaceae), P.I. Forster
(Asclepiadaceae), O. Gideon (Costaceae), N.H.S. Howcroft (Orchidaceae),
J. Pipoly (Myrsinaceae), P.F. Stevens (Clusiaceae, Ericaceae, passim), D.E.
Symon (Solanaceae), and G. Weiblen (Moraceae).

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VASCULAR PLANTS ON A GYPSUM OUTCROP IN
SOUTHERN NEW MEXICO: A LISTING, A NEW
VARIETY AND TAXONOMIC REALIGNMENTS IN
THE ANULOCAULIS LEIOSOLENUS COMPLEX
(NYCTAGINACEAE), AND A NEW VARIETY OF
MENTZELIA HUMILIS (LOASACEAE)

RICHARD SPELLENBERG and THOMAS WOOTTEN
Department of Biology
New Mexico State University
Las Cruces, NM 88003-8001, U.S.A.
ABSTRACT

A survey of the flora on a hitherto unexplored gypsum outcrop on the west face of the
Guadalupe Mountains in southern New Mexico has revealed a new variety of Anulocaulis
(Nyctaginaceae), A. /eiosolenus (Torr.) Standl.var. howardti ne & Wootten (var. nov.).

This variety has a combination of characteristics in various degrees intermediate to other
taxa, an observation that results in the inclusion of A. oni venus Watert. into A. leiosolenus
as A. /etosolenus var. i us (Watert.) Spellenb. & Wootten (comb. nov.). Also from this

outc ae is described Mentzelia humilis (A. Gray) J. Darl. var. gvadalupensis Spellenb. (ve
nov.). Fifty-nine species in 29 families were observed to occur on the gypsum outcrop. A
key is provided to all known Anw/locau/is taxa and a list is presented documenting the other
plant species that occur with it and the new variety of M. humilis.

RESUMEN

La ee de la nes en un afloramiento de yeso, inexplorado previamente, en la
adera oeste de las Montafi uadalupe en el sur de Nuevo México ha revelado una variedad
nueva de Ae (Nyc pe naeiey A. leiosolenus (Torr. ) Standl. var. howardti Spellenb. &
Wootten (var. nov.). Esta variedad tiene binaci6n de caracteres intermedios en varios

grados con otros taxa, una observaci6n que a como resultado la inclusion de A. gypsogenus

aterf. en A. /erosolenus asi como A. Jeiosolenus var. gypsogenus (Waterf.) Spellenb. & Wootten

(comb. nov.). También, Mentzelia humilis (A. Gray) J. Darl. var. guadalapensis Spellenb. (var.

nov. om esta ing de este etcramnient: Ci incaeniy nueve especies de 29 familias de sae
|

| ». Se ofrece una clave pa todos los taxa ¢

|

ye
de Alias y se presenta una lista ea

ylantas que conviven
con las dos variedades nuevas.

—_

INTRODUCTION
While deer hunting in southern Otero County, New Mexico, about a decade
ago, Michael Howard of the Bureau of Land Management, Las Cruces Dis-
trict, traversed a gypsum outcrop on the western slope of the northern por-
tion of the Guadalupe Mountains and noted an Anw/ocaulis. In the summer
of 1996 he showed the plant to Wootten, who collected a voucher, identified

Sipa 18(4): 987-999. 1999

988 Sipa 1L8(A4)

the plant as Anwlocaulis gypsogenus Waterf., and brought it to the New Mexico
State University herbarium for deposition. Spellenberg immediately rec-
ognized the plant as near A. gypsogenis but somewhat different. That col-
lection also brought attention to a previously unbotanized area on the west
face of the Guadalupe Mountains.

Wootten and Howard visited the southern portion of the gypsum out-
crop in September and November, 1996, and escorted Spellenberg to that
site in early August 1997. Spellenberg and Wootten returned to the south-
ern portion of the outcrop in September 1997 and April 1998, and to the
northern, more lengthy portion of the outcrop in July, 1998. Our survey
re-vealed new varieties of Anilocailis letosolenus and Mentzelia humilus, herein
described, and resulted in a collection of plants documenting the flora of
this gypsum outcrop (Appendix 1).

CHARACTERISTICS AND FLORA OF GYPSUM OUTCROP

This gypsum outcrop is part of the Permian age Yeso Formation (Hunt
1977). Most of the outcrop consists of limestone rubble, the stones 2—10
cm in diameter, or larger, overlying a pale tan, sandy, gypseous clay. In a few
places, when walked upon, t

ee

ye gypsum makes the hollow sound character-
istic of other gypsum outcrops in the Southwest. On the steeper slopes there
is little limestone rubble, the gypseous matrix being completely exposed.

The outcrop is not continuously exposed and consists of two main sec-
tions. The southern section extends for about 5 km ina NNW - SSE direc-
tion, and perhaps up to 3/4 km in an E-W direction, with an elevational
range of about 250 meters beginning at about 1350 m. It occurs on the
lower slopes of the western escarpment of the Guadalupe Mountains, be-
neath The Rim, at the upper edge of the bajadas that extend westward to
Crow Flats. The southern end of this section lies about 48 air km NNE of
Dell City, Texas, centered on about 32°22'N, 105°04'W (Sec. 25 to esti-
mated Sec. 11, T22S, RISE), just north of the mouth of Pup Canyon. Most
of the outcrop has a very steep western exposure, the slopes 30—45° and
steeper. Near the base of the outcrop at the southern end of the outcrop,
and also about 2 km to the south, there are a few low, gypseous hills chat
have slopes of all exposures, and within the major portion of the outcrop,
gullys and arroyos provide northern and southern exposures

The southern portion of the outcrop disappears near the northern end of a
small south-facing box canyon. It or a similar gypseous outcrop reappears on
the western slopes of the mountains about 3 km to the NNW. From there
the outcrop continues in a northwesterly direction in a discontinuous manner
for about 12 km (Sec. 33, T21S, RISE northward to Sec. 25, T20S, R17E).

This outcrop occurs in an remote area where access is comparatively dif-
ficult. The entire outcrop lies on public lands, some portions entirely on

SPELLENBERG AND WoorTeN, Vascular plants in southern New Mexico 989

those of the Bureau of Land Management, Las Cruces District, in other ar-
eas the eastern portions on the Guadalupe Ranger District of the Lincoln
National Forest. Browsing of a few of the species that occur on this out-
crop, probably by deer and rabbits, was noted; no sign of domestic live-
stock, common in the valley below, was observed. According toa local rancher,
Mr. George Rauch, the livestock simply do not get up to this rather inhos-
pitable site (he was speaking of the southern section, but conditions are
similar along the length of the outcrop). The two new endemics discovered
here presently do not seem to be impacted by human-related activities.

The Anwlocaulis and the Mentzelia are both common and conspicuous on
the outcrop. Because there are no records in collections of these taxa from
here, we believe that the outcrop had not been explored by botanists prior
to our work. Only a minor portion of the outcrop consists of highly con-
centrated gypsum, and here obligate gypsophiles such as A. /etosolenus, M.
humilus, and Selinocarpus lanceolatus occur. On this portion we document 59
plant species in 29 families (Appendix 1), including the two new varieties
described below. We stopped noting species when we crossed off the gyp-
sum and onto the cobbly limestone bajadas that support Chihuahuan Desert
vegetation consisting of Larrea tridentata (DC.) Cov., Parthenium incanum
Kunth, Acacia neovernicosa Isely, Fouquierta splendens Pagelay. Yucca torreyi
Shafer, Ephedra aspera Engelm., Dasylirion wheeleri 8. Wats., Echinocerenus pectinatus
(Scheidw.) Engelm., Krameria erecta Willd. ex Schinlees, Allionia incarnata
L., Stenandrium barbatum Torr. & A. Gray, Polygala macradenia A. Gray, among
numerous other species.

A NEW VARIETY AND TAXONOMIC REALIGNMENTS IN ANULOCAULIS

Spellenberg (1993) provided a taxonomic review of Anu/ocaulis and rec-
ognized five species, one of them, A. /ezosolenus, with two varieties. Turner
(1993) described a sixth species. This present paper reduces one of the spe-
cies recognized by Spellenberg toa variety in A. /e‘osolenus, and adds a fourth
variety to that species. We now consider Anw/locaulis to have five species,
one with four varieties.
aaa leiosolenus oe ) Standl. var. howardii Spellenb. & T. Wootten,

var. Type: UNITED STATES. New Mexico. Otero Co.: 48 km NNE of Dell
4 W base . Gos Mts., N of mouth of a Canyon, ca. 50 m SW of
26, T228, RISW, 32°22.14'N, 105°03.92'W, elev. 1360 m, 2a 997,
ee Wootten, and Howard 12433 (HoLoTYPE: NMC; isotypes: NY, TEX, UNM).
hAroae cpa Perer

A Anulocaule leiosoleno var. leiosoleno perianthiis meee roseis SoG albis vel s

foliis leviter glaucis et parce tuberculatis (vs. viridis non glauci | tuberculatis) differc.

Strong perennial from gnarled woody root. Stems 1-ca. 6, ascending, branched
in upper 70%, up toca. 1.2 m tall, glabrous, glaucous. Leaves usually in 2—
3 pairs in basal 1/4 of plant; petioles 35-50 mm long, blades more or less

990 Sipa 18(4)
orbicular, commonly wider than long, 45-105 mm long, 45-135 mm wide,
semi-glaucous, bluish-green, with sparse purplish pustules, pustules slightly
denser on abaxial surface. Inflorescences widely paniculate, forming the up-
per 2/3 of the plant, the flowers borne terminally and on short side branches
in more or less congested clusters. Buds with minute hairs at the apex.
Perianth deep rose-pink, obliquely funnelform, 22—32 mm long, limb 10—
17 mm wide, stamens exserted ca. 20 mm, the style 25 mm. Fruit biturbinate,
4.3-5.9 mm long, 3.7—4.5 mm wide, with 10 irregular longitudinal ridges
and an equatorial wing 0.2-0.9 mm wide.

At present, the new Avw/ocaulis is known only from the western slope of
the Guadalupe Mountains in south-central New Mexico on tannish, shaley,
gypseous clays, and then only from the southern portion of the gypsum outcrop,
immediately north of Pup Canyon. Exploration of the outcrop to the north
revealed no other populations. We name the plant for its discoverer, Michael
Howard, of the Las Cruces District of the Bureau of Land Management,
whose attention to natural biota resulted in the discovery of this Anilocanlis.
Mike has a strong sense of responsibility for the nation’s natural resources,
shares his knowledge willingly with the public, and works toward making
land-use in the Southwest compatible with needs for conservation.

Paratypes: NEW MEXICO. Otero Co.: ca. 30 air mi NE of Dell City, Texas, foothills
of Guadalupe Mtns. at mouth of Pup Canyon, extreme SE corner of Sec 23, T225, RISE,
26 Sep 1996, Wootten and Howard s.n. (NMC); 48 km NNE of Dell City, Texas, W base of
Guadalupe Mts., N of mouth of Pup Canyon, edge of Lincoln National Forest, ca. 300 m
ENE of SW corner $24 T22S R1I8W, 32°22.22'N, 105°74'W, elev. 1360 m, 5 Aug 1997,
Spellenberg, Wootten and Howard 12435 (ARIZ, NMC, UC); 49 km NNE of Dell City, Texas,
W base of Guadalupe Mts., N of mouth of Pup Canyon, NE Sec 23, T22S, RI8W, 32°22.75'N,

105°03.99'W, elev 1460 m, 5 Aug 1997, Spellenberg, Wootten and Howard 12438 (NMC).

Anulocaults letosolenus var. howardii is immediately distinguishable from
close relatives by the combination of the semi-glaucous leaves with only a
few large multicellular trichomes, the reddish-pink perianth, and the moderately
broad rim on the fruit (Table |). The fruit is most similar to that illustrated
in Spellenberg (1993, Fig. LF), a fruit from a Texas race of A. /esosolenus var:
leiosolenus, Within the complex, minute hairs at the tip of che perianth, best
seen in bud, were previously known only in the var. lasianthus.

As illustrated in Table 1, the new variety has some of the characteristics
that are used alone or in combination to distinguish among other members
of the A. gypsogenas-leiosolenus complex (Spellenberg 1993). For this reason
we believe the newly discovered entity is best recognized as a variety in a
more broadly reconstructed A. /esosolenus, in which A. gypsogenus is included
at the varietal level. The entire complex presents a classic representation of
completely allopatric, closely related, more or less distinguishable races, this
structure commented upon by Spellenberg (1993). Mayr (1969, ch. 3) dis-

SPELLENBERG AND WoOorTTEN, Vascular plants in southern New Mexico pepe

TABLE 1. A comparison of some characteristics used to distinguish varieties
within Anwlocau/lts letosolenus.

Variety of A. Perianth limb Perianth Leaves Fruit wing
leiosolenus color pubescence
LySpsogenus white to very none pale bluish green, 0.8—1.2 mm
pale pinkish glaucous, smooth,
pale gray or whitish
when driec
howardii deep rose pink minute trichomes _ bluish green, 0.2-0.9 mm
at tip in buc semi-glaucous,
few largeconical
tubercules, grayish
green when dried
lasianthus pale pink to pink ~=minute trichomes green, semi-glaucous,
at tip in bu dense conical ).2-0.6 mm
tubercules, dull
green when dried
lerosolenus white to pale pink none green, not especially 0.2—0.6 mm

glaucous, dense conical
tubercules, dull green
when dried

cussed problems with “eciding taxonomic divisions in allopatric popula-
tions; Stuessy (1990, ch. 12) discussed problems with assigning infraspe-
cific taxa to varieties or subspecies. With regard to Anu/ocaulis, the tradition
of using variety is followed for taxonomic recognition of closely related,
internally rather homogeneous populations that can be distinguished from
other, similar populations.

Spellenberg (1993) noted the similarity between the large, pale perianth
of the western race of A. /eiosolenus var. leiosolenus and that of A. gypsogenus.
Ona more subtle note, the var. howardii has flowers clustered in the inflo-
rescence reminiscent of var. /aszanthus. This characteristic is somewhat de-
veloped in A. gypsogenus and much less so in the var. /ezosolenus. The leaves of
the var. howardii are more similar to A. gypsogenus. The fruits are rather in-
termediate between A. /ezosolenus and A. gypsogenus, as traditionally recog-
nized. It is our view that with the discovery of the easily recognizable race
now named as A. /ezosolenis var. howardii, the other phases in this complex
of gypsophilic endemics are best recognized as belonging to one geographically
fragmented (Fig. 1) species of variably differentiated allopatric races. For
that reason, we transfer A. gypsogenus into A. leiosolenus as a variety restricted
to the gypsum along the Pecos River, slightly to the east of, but completely
disjunct from var. howardii.

992

Sipa 18(4)

* NEVADA

CALIFORNIA

BAJA
CALIFORNIA

C1 var. gypsogenus = SONORA
@ var. howardii

© var. leiosolenus

© var. lasianthus

“i,

of states of southwestern United States and northwestern Mexico sh
sisecioution of Anilocaulis leiosolenus varieties.

CHIHUAHUA

Fic

=

. Map

By

Anulocaulis eechegaioue (Torr.) Standl. var. gypsogenus (Waterf.) Spellenb.
a Wootte n Ov. i ASIONYM: Anulocaulis eypsogenus Waterf.
329. oe Type: L et iD STATES. New Mexico, Chaves Co.: C Baienclean Blufts,

7 oe E of Roswell, 9 Oct a a 5701 (HOLOTYPE: GH!; lsorypes: NY! MO!).

om

, Rhodora

KEY TO TAXA OF ANULOCAULIS
1. Ancthocarp without a prominent equatorial ridge or wing; perianth less than
16 mm long, the lobes flaring but not reflexed.
2. Perianth 15-16 mm long at anthesis, purplish, che cube externally gla-
brous, minutely glandular-pubescent near the apex; anthocarp ellipsoid
or broadly fusiform (southern Coahuila). 0.0.0.0... A. hintoniorum B. L. Turner
2. Perianth less than 10 mm long at anthesis, whitish to rose-lavender, the
tube externally villous; ancthocarp fusiform or turbinat
. Flowers usually 5-15 in umbel-like clusters; cube of psa not elongating
after anthesis: anthocarp broadly fusiform (southeastern California).

—
eae

A. earns (Coville) Standl.
. Flowers usually borne singly; tube of perianth markedly elongating after

ae anthocarp curbinate, bluntly 5- sae ridges indefinite peutliem
Texas, western Coahuila, possibly eastern Chihuahua). ................ . eriosolenus
(A. Gray) Seandl.
|. Ancthocarp with a prominent equatorial ridge or wing; pertanth 10-35 mm
long, the lobes flaring or sharply reflexed

SPELLENBERG AND WoorTTeN, Vascular plants in southern New Mexico 993

4. Perianth ca. 10 mm long, the lobes sharply reflexed (northeastern Chi-
huahua and immediately adjacent Texas). A. reflexus I. M. Johnst.
4. Perianth 22—35 cm long, the lobes flaring, not reflexed «0.0.0.0... A. leiosolenus
(Torr.) Standl.

5. Leaves smooth, glaucous; perianth glabrous externally at apex (Pecos

River in southestern New Mexico and western Texas). .........008 var. gypsogenus
Waterf.) Spellenb. & T. Wootten

5. Leaves at least sparsely tuberculate; perianth minutely puberulent or

glabrous at the apex
6. ieee glabrous emly at the apex (extreme western Texas, south-
al New Mexico, north-central Arizona, and southern Nevada)
var. leiosolenus

6. Perianth minutely pubescent externally at the apex (visible best when
in late bud).
. Leaves grayish green; purplish tubercules on leaves sparse; peri-
anth deep rose-pink (south-central New Mexico, W face of Guadalupe
ar. howardii Ba ae & T. Wootten
eaves green or dark green; purplish ae on leaves dens
perianth pale pink to pink (Big Bend region of Texas and imme
diately adjacent Chihuahua) ar. lasianthus I.M. Johnston

~I

~~

A NEW VARIETY OF MENTZELIA HUMILIS

Mentzelia humilis (Urb. & Gilg) J. Darl. is a variable yet distinctive member
of sect. Bartonia Torr. & A. Gray restricted to gypseous substrates in south-
eastern New Mexico and western Texas (Thompson 1997). Leaves vary from
pectinate to entire. Usually leaves in a population are similar, but occasion-
ally populations will have pectinate and entire leaves (Sivinski and Lightfoot
2634; cited Mentzelia specimens comprise Appendix 2). Leaves may be clearly
pectinate, the lobes distantly spaced and linear, 1-2 mm wide and about
15 mm long, or lobes may be much shorter. Lobes may be straight or some-
what falcate, curving toward the leaf apex. In either case, the margins of a
lobe are approximately parallel and the apex of the lobe is almost always
rounded. In some cases lobes are completely absent and leaves are entire
and linear. Basal leaves may be much less lobed than cauline leaves (Hzggins
6845). Other than by flower color (not given, but presumably white) H7ggns
6845, from western Texas, cannot be distinguished from a pectinately lobed
form of M. perennis H. J. Thompson (ined.) from central New Mexico (Edwards
and Repass 47206; Spellenberg and Willson 4233; Ward et al, 81-281), leaving
one to ponder the distinction of these two taxa.

Specimens of this complex in NMC have been annotated as either M.
humilis or M. perennis by H. J. Thompson. Martin and Hutchins (1981) separated
these two taxa in their key on a vegetative character, tufted (ML. perennis) vs.
not tufted (M. Awmilis), a feature that will not distinguish them. They give
flower color of the former as “pale lemon-yellow,” which is approximately
correct, vs. “yellow” for M. humilis, which ts incorrect (pale ochroleucuous

994 Sipa 18(4)

to white). Perhaps Martin and Hutchins were following to some degree Wooton
and Standly (1915) who noted petals to be “pale yellow” (in Nuttallia gypsea
Wooton & Standl., a synonym of M. Awmilus), or Darlington (1934), who
“keyed” M. humilis under “flowers lemon-yellow to golden.” This assump-
tion of yellow-colored flowers is pect aar able because buds are cream and

dried petals in fresh specimens are definitely yellowish. Thompson and Zavortink
(1970) may be the first to have indicated that M. Awmz/is corollas and androecia
were white. Later Thompson (1997) described the petals as “white or very
pale yellow.” Mentzelza perennis has pale yellow petals.

In Thompson (1997) and Thompson and Zavortink (1970), Mentzelia
specimens from the gypsum outcrop discussed earlier key to M. strictissima
(Wooton & Stand.) J. Darl., but this is a very different, tall plant chat oc-
curs in (often) sandy soil. It has dentate leaves and cylindrical capsules. Leaves
of the novel Mentzel7a much more closely resemble those of M. mexicana H.
J. Thomps. & Zavort. or M. saxicola H. J. Thomps. & Zavort. as illustrated
in Thompson and Powell (1981, fig. 7). These are yellow-flowered species
of western Texas and northern Mexico once confused with M. sz/tiflora (Nutt.)
A. Gray. Unfortunately, Thompson and Powell did not discuss the relationship
of M. humilis to any of these species. In a survey of other specimens, leaves
from the novel plants from the gypsum outcrop on the western slope of the
Guadalupe Mountains more closely resemble those of some specimens of
M. multiflora, and also resemble that illustrated in Thompson and Powell
(1981, fig. 7) for this species.

Thompson and Powell (1981) illustrated, described, and compared the
seed coats of M. multiflora with seed coats of M. mexniuuva and M. saxicola.
They noted the first to have cells with swollen outer walls covered by nu-
merous small papillae. This gives the seed coat a coarsely granular appear-
ance or, as stated in Thompson (1997), it is “rough with papillae. ” Seed
coats of the latter two have ae papillae and appear “smooth” (Thompson

1997), but actually are very finely granular under a microscope at about
20x. Thompson did not give the characteristics of seed coats of M. humilis;
we note them to be very similar to those of M. mv/tiflora, as are the seed

—
—

—

coats of the novel Mentze/ia in question.

Thompson, in attempting to work out distinctions between New Mexico
populations of M. mu/tiflora, M. jemezensis, M. humilis, and M. strictissima
sent Spellenberg a letter (29 Apr 1980), a map, and color photographs (without
provenance) explaining his interpretation (filed at NMC, accession #605 36,
in M. jemezensis folder). Mentzelta multiflora and M. humilis have very dis-
tinct flowers, the first yellow (Thompson’s photo is more yellow than most
races in southern New Mexico, which may be pale yellow), M. Awmilis near
white. More important M. mu/tiflora has broader petals with the transition
to stamens with expanded filaments comparatively abrupt, whereas M. Awmilis

SPELLENBERG AND WoorTTeN, Vascular plants in southern New Mexico 02)

has narrower petals, the transition to stamens more gradual. Flowers from
the new Mentze/ia in question very strongly resemble those of the photo
and of specimens of M. humilis and are not like those of M. mu/tiflora.

Thompson (1997) indicated capsules of M. multiflora to be cylindrical,
15—25 mm long, whereas he wrote that M. humilis has capsules cup-like,
6-13 mm long. The novel Mentze/ia has capsules that are cup-shaped and
in the lower range of length for those of M. Aumailis. Plants of the new popu-
lation are densely clumped, like some races of M. humilis, and leaves are
sub-entire, dentate, or pinnatisect, reminiscent of those of M. mu/tiflora. When
the leaves are pinnatisect, the lobes taper from a broad base to a narrow,
acute or even acuminate tip. The flowers and capsules are like those of M.
humilis. The inflorescence is much more congested than in any of the spe-
cies mentioned.

The map Thompson provided with his letter to Spellenberg (29 Apr 1980)
shows M. Aumilis to occur in western Texas and eastern New Mexico in the
Pecos River drainage, distinctly, but not distantly, east of the population in
question. He maintained this distribution for M. Aumi/7s in his 1997 manuscript.
On his map accompanying the letter, M. perennis is shown to occur ina lim-
ited area to the northwest of the site from which the new Menfzelia origi-
nates. Mentzelia multiflora, in contrast, is widespread in the western United
States and northern Mexico (Thompson 1997). It is known from robust to
smaller plants in the Guadalupe Mountains (¢.¢., Spellenberg 3660, Wooton
s.n.). Both these specimens have broad petals and cylindrical capsules rep-
resentative of the species; Spellenberg noted flower color as “pale yellow”
on the specimen label. In his letter Thompson alluded to the possibility of
gene flow between isolated edaphic endemics and more widespread edaphically
unrestricted species. Such a process might explain in the new variety the
leaves similar to M. multiflora and the flowers and capsules similar to M.
humilis. Observations from these populations reveal that flowers and cap-
sules of the novel Mentzelia are consistent, foliage and habit are variable.
Nevertheless, the race is consistently distinct from M. Aumalis var. humilis
and is geographically isolated from it: var. humilis east of the Guadalupe
Mountains, var. gvadalupensis restricted to the western slope.

Mentzelia humilus (A. el, ae Nee Steuer Seams var. Nov.

(Fig. 2).Typs: UNITED S Mexico. Otero Co.: 48 km NNE of Dell

City, Texas, W base Gun ae N eet ith of Pup Canyon, $14 T22S R18W,

32°22.74'N, 105°04.26'W, elev. 1460 m., 22 Sep 1997, Spellenberg & Wootten 12455
(HOLOTYPE: NMC; tsorypes: NY, TEX, UC, UNM).

Mentzeliae humili var. humili foliis dentatis vel pinnatisectis (vs. pectinatis vel integris),

rachidibus 2—8 mm (vs. 1—2 mm) latibus, inflorescentiis congestibus (vs. noncongestibus),

et plerumque pedicellis capsulis iievieubus (vs. aequantibus vel longioribus) differt.

Plants 0.5—2.5 dm tall; basal leaves spatulate, dentate, with 2—4 teeth

996 SipA 18(4)

2. Illustration of plants comprising the type collection of Mentzelia humilis var. guadalupensis
Ce: & Wootten 12455). Collection was made with the intent of illustrating the ma-
jor aspects of variation in the population: habit and leaf dissection. All plants are clearly

All plants in photograph

—

identified in distributed specimens by small tags affixed to then
are at NMC. Plant A is the holotype plant, which has been aes separately from isotypes

t NMC; other portions of plant A have been distributed with other isotype plants to NY,
UC; plant B, NMC, NY; plane C, NMC, NY; plant D, NMC, TEX, UC, UNM. Plants
similar to A—C are common in the population; plants with leaves dissected to the extent of

plane D are less common.

SPELLENBERG AND WooTTEN, Vascular plants in southern New Mexico 91

per side, or sometimes the smallest basal leaves nearly entire; midstem leaves,
3.0-8.5 cm long, 8-21 mm wide, the blade spatulate or lanceolate in out-
line, tapering gradually to a slender petiole, the blade from shallowly to
deeply dentate or pinnatisect, 3—7 teeth or lobes per side, when pinnatisect
or deeply dentate the rachis 2-8 mm wide and the lobes straight or some-
times falcate, tapering from base to acute tip; bracts beneath the heads pinnatisect,

—4 lobes per side; petals white or pale ochroleucous when fresh (drying
ochroleucous), about 10 (intergrading with the outer stamens with broad
filaments), 1O-13 mm long, 1.6—2.1 mm wide, acute; capsules cupulate,
5-8 mm long, 5—6 mm wide, the length 1—1.7 times the width, the calyx
lobes on capsules 4-9 mm long, narrowly triangular-subulate; seeds 1.8—
2.2 mm long witha wing 0.5 mm wide, the seed coat conspicuously papil-
late (use 20x), the papillae hemispheric and minutely granular.

As far as is known, Mentzelia humilis var. guadalupensis is restricted to the
west slope of the Guadalupe Mountains on gypsum of the Yeso Formation,
probably occurring throughtout the outcrop as described in the introduc-
tion. The varietal epithet refers to its presence in the Guadalupe Mountains.

ParatyPEs: NEW MEXICO. Otero Co.: 49 km NNE of Dell City, Texas, W base of
Guadalupe Mcs., N of mouth of Pup Canyon, NE $23, T22S, R18 W, 32°22.75'N, 105°03.99'W,

elev. 1460 m, 5 Aug 1997, Gillon "Wootten, and Howard 12440 (NMC); 2.5 km S of the
Chaves Co. line, oa west oe Rim on the west se of the Guadalupe Mts., about 100
m inside the Lincoln National Forest, center $7, T21S, RISE, 32°29.65'N, 105°08.28'W,
elev. 1540 m, a Jul 1998, pene G& Wootten pe (NMC, RM).

APPENDIX |
The following list provides the names of plants that we collected or ob-
served on the outcrop; collection numbers are Spellenberg’s. Deposition of
specimens is indicated by herbaria codes as presented in Holmgren et al.
(1990). Each name is also followed by “C, O, or U,” signifying generally
“common, occasional, or uncommon,” respectively, on the outcrop.

Agavaceae: Dasylirion wheeleri S. Wats. {O}; Yucca elata Englem. {U}; Yucca torreyi Shafer [O}
Amaranthaceae: Tidestromia suffruticosa (Yorr.) Stand. var. suffruticosa, 12481 (NMC) [U}
Anacardiaceae: Rou microphylla Engelm. {U}
Apocynaceae: Amsonia longiflora Torr. var. salpignatha (Woodson) McLaughlin, 12434 (NMC,
NY, U ), 12441 (NMC ), 12474 (NMC, NY), 12501 (NMC, NY) [C]
Asclepiadaceae: Asclepias macrotis Torr., 12507 (NMC, NY) {U]
Asteraceae: Brickellia laciniata A. Gray {U]; Gaillardia multiceps Greene [U}]; lle reZld
Mis ee ) A. Gray [O}; Haploisthes greggii A. Gray, 12437 (NMC) {C}; Mac
mnatifida (Hook.) Shinners var. pinnatifida, 12505 (NMC) [U}; Porophyllum scoparinm
i“ Gs 12436 (NMC, UC) [C]; Sartwellia flaveriae A. Gray {U]; Thelesperma megapotamicum
(Spreng.) Kuntze {U]; Thymophylla acerosa (DC.) Strother, 12484 (NMC) {U]; Thymophylla
pentachaeta (DC.) Small var. hartwegii (A. Gray) Strother, 12475 (NMC, NY), 12483 (BRIT)
Rial stenoloba 8. F. Blake [O
Boeoiia ceae: Tiguilia hispidissima (Torr.) A. Richardson, 12502 (NMC) [O]
Brassicaceae: ees camporum (A. Gray) Greene, 12442 (NMC) [C}

’ > =

998 Stipa 18(4)

Cactaceae: C oe ee E ange Im.) A. bee 12482 (NMC) [U}, Echinocactus
horizonthatonins Lemaire {U}; E us Englem., 12486 (NMC) [O}; Opantia
imbricata Haw.) DC, a oe macrocentra Engelm., 12487, spineless (NMC), /2488,
spines (NMC) [O]; Opuntia phaeacantha Englem. var. phaeacantha {U]}

Chenopodiaceae: Atriplex canescens <> Nutt. {U i

Ephedraceae: Ephedra aspera 8. Wats., 12473 (NMC) {O]

Euphorbiaceae: Chamaesyce fendleri ( an & Gray) Small. 12458 (NMC), 12471 (NY) {O],
Croton dioicus Cav. {O}

Fabaceae: Acacia neovernicosa Isely

Fouquieriaceae: Mouguieria ieee ae ‘lm. [O}]

Hydrophyllaceae: Nama carnosum C. L. Hitche., 12503 (NMC), [O]

Kramertaceae: Krameria erecta Schult., (12478) {O]

Lamiaceae: Hedeoma nanum (Torr.) Brig., 12479 (NMC) [U}

Linaceae: Linum vernale Wooton, 12480 (NMC) {U}

Loasaceae: Cevallia sinuata Lag. {U}; Mentzelia te (A. Gray) J. Darl. var. ee
Spellenb., 12440 (NMC), 12455 (NMC, NY, 7 UNM),/2500 (NMC,

Malvaceae: Sphaeralcea coccinea (Nutt.) Rydb., / oe - (NMC ){U]

Nyctaginaceae: A//ionia incarnata L. var. incarnata (O}, Anilocantis lerosolenns (Torr.) Stand).
var. howardii Spellenb. & T. Wootten, 12433, 12435, 12438 [C]}; Mirabilis linearis (Pursh
Heimerl, 1/2506 (NMC) {U}; Sol nonarp lanceolatus Wooton var. alae. 12452(NMC),
12499 (NMC) [O

Oleaceae: Menodora scabra A. Gray {U]

Onagraceae: Gaura coccinea Nutt. [U}; ve lophus hartwegii (Benth.) Raven subsp. filifolias
(Eastw.) Towner & Raven, 1/2472 (N .

Poaceae: sa sh Nutt. var. ie Vasey) Allred, 12453 (NMC) [U]; Aristida
pansa Wooton indl. var. pansa, ee {U}; Bothriochloa laburoides (DC.) Herter
ee torveyana Seeud ) Allred & Gould, 12459 (NMC) {U}; Bowteloua Niue Gould

adia, 12443 (NMC) [O}; Dasyochloa pulchelta ( (Kunc.) Steud. [U}; Diestaria cognata
. ae ) Pilg. sabes i Wiptf& Hatch, 12457 (NMC) {U]; Setaria lencopila (Scribn.
& Merr.) K. Schum. [U}; Sporobolus cryptandrus (Torr.) A. Gray, 12504 (NMC) [O}; Stipa
curvifolta a len, / 2477 (NMC, NY) {U}, Virrdens maticus (Yorr.) Nash var. muticus, 12454
(NMC) [U

hia eae Ertogonum havardit 8. Wats., 12439 (NMC) {C}
teridaceae: Astrolepis cochisensis (Goodd.) D. M. Benham & Windham, 12444 (NMC) [O]

. Pallugia pavadoxa (D. mee Endl. [U

Rubiaceae: Hedyotis nigricans (Lam.) Fosberg, 12445 [O}

Solanaceae: Nicotiana ene Dunal (NMC) {U}

—

APPENDIX 2
Collections of Mentze/ia cited in discussion of M. humilus var. guadalupensis.
eee of specimens is indicated by herbaria codes as presented in Holmgren
et al. (1990).
ai zelra humilis (A, Gray) Darl.—H gers 0845, Texas, Culberson Co., 26 mi E of Fey.
-180 along Hwy 652, gypsum soil, 21 May 1973 (NMC); Stvinski & Lightfoot 2634
New Mexico, Guadalupe Co., 2.3 mi S of Pecos ] bridge at Puerco de Luna, 21 oe tC
1993 (NMC, UNM)
Mentzelia multiflora (Nutt.) A. Gray.—Spellenberg 3660, New Mexico, Otero Co., Guadalupe
Mts. on Guadalupe Rim Rd #67, 8 Sep 1973 (NMC), (Mis sa., New Mexico, [with-
out county}, Guadalupe Mts., wese slope, 3 Aug 1909 (NMC

SPELLENBERG AND WoorTTeN, Vascular plants in southern New Mexico 999

Mentzelia perennis Wooton.—Edwards & Repass 4726, New Mexico, Socorro Co., ca. 8 mi E
of Socorro, 23 Jul 1977 (NMC); eine & Willson 4233, New Mexico, Socorro Co., 8
mi (by air) ENE of Bingham, W edge Chupadera Mesa, 8 Jul 1976 ee Ward, pp
& Soreng 81-281, New Mexico, Lincoln Co., W base of Cerro Tecolote SSW
of Corona, roadside or US 54, 3 Jul 1981 (MO, NMC, eee. ienbhes as M.
pumila (Nutt.) Torr. & A. Gray).

ACKNOWLEDGMENTS

We thank several individuals for assisting in various ways with this study.
Mike Howard brought the Avw/ocau/is to our attention and accompanied
us to the site on our first visits. Jane Shafer and George Rauch, ranchers on
Crow Flats in southern New Mexico, allowed access to the site across deeded
land. Manual Tanner and J.T. Tanner allowed access from their leased land
to the gypsum outcrop to the north of the Pup Canyon outcrop. Photo-
copies of specimens of Mentzelia mexicana from SRSC were supplied by A.
Michael Powell. Richard Worthington supplied information from UTEP.
Help with Latin diagnoses was received from John Strother and Alan Smith,
and Strother carefully read a preliminary version of the manuscript. Robert
Kiger assisted with a nomenclatural problem. A thoughtful and very help-
ful review of the manuscript was provided by Denis Kearns.

REFERENCES

DarLiNGton, J. 1934. A monograph of the genus Mentze/ia. Ann. Missouri Bot. Gard. 21:103—
22

Ho iMGren, P.K., N.H. HotmMGren, and L.C. BARNETT. 1990. Index Herbariorum, Part I:
The Herbaria a the World, 8th ed. Regnum Veg. 120:1-693.

Hunt, C. 977. Surficial geology of southeast New Mexico. Map GM-41. N. M. Bureau
of anes N. M. Institute of Mining and Technology. Socorro, NM

Martin, W.C., and C.R. Hutrcuins. ee A flora of New ae vol. 2. J. Cramer, Vaduz.

Mayr, E. 1969. Principles of systematic zoology. McGra iL; Ines IN

SPELLENBERG, R. 1993. Taxonomy of pera aan Sid 15:373-389.

Srugssy, T.F. 1990. Plant ee Columbia University Press

THOMPSON, H.J. 1997. Loasaceae. In: J. Henrickson and M.C. idhacian, rong a of the Chihuahuan
Desert Region, draft edition 1.2, Vol. 1, published by J. Henrickson, Los Angeles, CA.
Pp. 513-520.

and A.M. Powe. 1981. Loasaceae of the Chihuahuan Desert Region. Phytologia

49:16-32.

and J.R. Zavortink. 1970. Loasaceae. In: D.S. Correll and M.C. Johnston.
Manual of the vascular plants of Texas, Texas Research Foundation, Renner, Texas. Pp
1082-1087.

Turner, B.L. 1993. A new species of Anu/ocaulis (Nyctaginaceae) from southern Coahuila,

Mex
ces E. O. and P.C. StanpbLey. 1915. Flora of New Mexico. Contr. U.S. Natl. Herb.
9:1-794.

1000 Sipa 18(4)
BOOK REVIEW

ANNE OrtH Epp te and Lewis E, ee 1995. A Field Guide to the Plants
of Arizona. (ISBN 1-56044-314-6, pbk.) LewAnn Publishing Co.,
Mesa, Arizona, with SkyHouse ee an imprint of Falcon Bick
Publishing Co., Helena, MT. Distributed by Falcon Press Publishing
Co., Inc., PO. Box 1718, Helena, MT 59624. $24.95. 347 pp., 899
color photos.

This ambitious field guide boasts almost 900 color photos picturing 850 of Arizona’s over

3,000 species of plants. It covers wildflowers, cacti, trees, shrubs, vines, and ferns. The

grasses and “weeds,” as is often the case, are left out. After a brief introduction on scope

and use, the six life zones of Arizona are delineated by as rainfall, and vegetation
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>

rally lic, fill flash, and p
by two photos: one habit shot and a close up, or fruiting shot etc

The photo section begins with the ferns and is followed by the trees, which are grouped
by leaf shape. The cacti, grouped by general pad morphology, then logically segue into the
agaves. Wildflowers and shrubs with conspicuous flowers are next, grouped by flower color,
number of segments (basically number of petals), and sometimes regular versus irregular
flowers. These and other additional sub-groupings make the photo section more diagnos-
tic and useful compared to the simply color grouped format found in many field guides.
Each photo ts sequentially numbe se followed by a common name, and referenced to a
text (

Next come the 295 pages of Tee ne Though some entries are somewhat brief all

sage where the description is fou

offer useful and interesting information. The text section is grouped phylogenetically by
family. Each family entry has a common name, the scientific name (with phonetic pronun-
ciation) and a general description of the family. Each species entry has one or more com-
mon names, the scientific name, description, elevation, habitat, and a comments section.
“he description sections vary in length and in some cases are somewhat spare but in other
cases are very ee and botanical, requiring some facility with technical terms . What
ever the descriptions may lack is more than made up by the comments section of each en-
try. | found them chocked full of ae little bits of information such as wildlife uses,
medicinal uses, toxicity, derivation of the plant names, and addition curiosities of che plane.
Also included is a pronunciation of the genus and the number of different species of that
genus found in Arizona, and finally the location and date of the photo.
rally like the glossary that follows. Besides containing the usual defined botanical

—

terms, it also includes the definitions of many Latin descriptors used in scientific plane
names along with their various endings (Bravo!). This is a real plus for the amateur at-
rata to tackle the mumbo jumbo of the Lace lingo. And, no, I wouldn’t leave out the
index, the real proof of the pudding. Thirty-two pages of wonderfully readable and un-
cluttered bold-faced genera and scientific ie names interspersed among the regular type
faced common names and specific epithet

This is a must have for anyone ene in the flora of Arizona. Practically every photo
is a masterpiece and there is plenty of information in the text to keep anyone interested.
Congratulations on a huge project so well-executed.—Robert J. George.

Sipa 18(4): LOOO. 1999

A NEW SPECIES OF PORTULACA (PORTULACACEAE)

JAMES HENRICKSON
Department of Biology
California State University
Los Angeles, CA 90032, U.S.A.

THOMAS R. VAN DEVENDER

Arizona-Sonora Desert Museum

Tucson, AZ 85743, U.S.A.

ABSTRACT

Portulaca yecorensis is described from the conglomeritic mudflow — at 1500-1600
m elevation in the Yécora valley in the Sierra Madre of eastern Sonora, Mexico. This an-
nual is related to P. oleracea and P. retusa but differs in small size, strong ene h pigmenta-
tion, smaller seeds, larger flowers and pollen, discribution, and habitat.

RESUMEN

Se describe la especie nueva Por re yecorensis del Valle de Yécora de la Sierra Madre
Occidental en la parte oriental de Sonora, México; crece en areas con conglomerados de
eee casi sin vegetacién. Esta especie anual esta relacionada con P. oleracea y P. retusa,

»p aaa rojiza fuerte, semillas mas pequefias,

Agia: y polen mas wena, digabucién, y habitat.

Key woros: Portulaca, Portulacaceae
INTRODUCTION

As part of their floristic surveys in the Sierra Madre Occidental of east-
ern Sonora, México, Tom Van Devender and Ana L. Reina G. encountered
a surprisingly rich flora on gentle hills and level areas at about 1500-1600
m elevation near Yécora, Sonora. The plants growing on barren, apparently
edaphically severe sites, are of particular interest. These surfaces are on de-
bris and conglomerate deposits in the Baucarit Formation that formed as
mudflows sometime after the beginning of the modern basin and range
morphology (17 million years ago, Cochemé & Demant 1991).

ith a mean annual rainfall of 913 mm/yr at Yécora (Burquez et al. 1992),
the vegetation is grassland in the valley bottom and pine-oak forest on the
surrounding slopes (Reina et al. 1999), Dominant trees in pine-oak forest
are Pinus engelmannii (Apache pine) and P. yecorensis (Yécora pine). Other
common forest trees include P. chihuahuana (Chihuahua pine), Quercus arizonica
(Arizona oak), Q. chihuahuensis (Chihuahua oak), Q. durifolia, Q. oblongifolia
(Mexican blue oak), Q. viminea (willowleaf oak), and Juniperus deppeana (tascate,

Sipa 18(4): 1001-1007. 1999

1002 Sipa 18(4)

alligator bark juniper). East of Yécora, shallow soils on mudflow surfaces
support open oak woodland-grassland transition with common Q. chihuahiensis
and Q. toumeyi (Toumey oak).

Considering the moderately high rainfall at Yécora, bare areas without
vegetation and minimal soil development on the mudflow surfaces are not
easily understood. Perhaps erosion rates are excessive. In the spring, the surfaces
are very dry with few visible plants. However, from July through Septem-
ber, heavy rainfall keeps the surfaces wet and a diverse dwarf herb flora flourishes
including sedges (10 species in 7 genera), succulents (Agave polianthiflora,
Echinocerens stoloniferus var. tayopensis, Mammillaria saboae var. handeana, M.
wrighti var. wilcoxit, Sedum vinicolor, Talinum marginatum), grasses (Microchloa
kunthit, Mublenbergia annua, M. shepherdit, M. texana), legumes (Aeschynomene
americana vat. glandulosa, Dalea confusa, D. filiformis, Stylosanthes sp.), and many
others. Thus far, four taxa have been described as new from the mudflow
areas including Mammuillaria saboae var. haudeana (A. Lau & Wagner) Glass &
R. Foster, Menodora yecorana T. Van Devender & B.L. Turner, Pectis vandevenderi
B.L. Turner, and Tridax yecorana B.L. Turner. A new Boerhavia known only
from these mudflow surfaces is under study by Richard Spellenberg and Luis
A. Pérez. To this list, we add a small, reddish, annual succulent Portslaca.

TAXONOMY

Portulaca yecorensis Henrickson & T.Van Devender, sp. nov. (Figs. I, 2).
Type: MEXICO. Sonora: Municipio de Yécora, 5.0 km NW of Yécora on ro to
Mesa Grande, sparse pine-oak ene. level mudflow deposits, 28°24'27"N, 108°57
1600 m, 29 Sep 1998, TF R. Van Devender 98-1942, with A.L. Reina G., W. rs
(HOLOTYPE: ARTZ; isorypes: CAS, TEX, MEXU).

>

=

Plantae annuae, rubro-marroniae; folta leviter compressae, 2—7.5 mm longae, 1.53.5
mm latae, petioli curtae; flores LO-20 mm diametro, petala aureae, fortiter emarginatae;
stamina |2—17; lobi styli

=

3-4; seminae 0.7—0.8 mm longae, parum compressae

Apple red to nearly maroon, succulent, low, spreading-ascending annu-
als, locally common on exposed shallow-soiled mudflow surfaces, the plants
(1—)2.5—5.0(—10.5) cm in diameter; stems 0.7—1.0 mm in diameter, alter-
nate or opposite branched, with internodes 0.5—8 mm long. Leaves fleshy,
the blades ovate to obovate, 2—7.5 mm long, 1.5—3.5 mm wide, slightly
compressed, obtuse to rounded at the tip, broadly obtuse-rounded at the
base above obscure petioles 0.2—0.7 mm long; axillary hairs very sparse,
obscure, to 0.3 mm long. Flowers 2—3(—4) at the stem tips where subtended
by clusters of sessile leaves; bracts 3, fleshy-membranous, ovate, reddish,
conduplicate, attached along a broad base, obtuse, the bracts 1—-1.8 mm
long, the paired bractlets 0.9—1.2 mm long, all persistent; pedicels ca. 0.5—
0.7 mm long, ca. 0.7 mm wide, expanding to 1.5—2.5 mm wide at the lower
hypanthial margin; sepals 2, apple red, glabrous, conduplicate with one external

HENRICKSON AND VAN DevVENDER, A new species of Portulaca 1003

ye Noe oR Phar 1a i) Ne eet as ts ie :
ah eS en < si Nipon wecee Cee Wanwig GB ty oS Oe fee ASR Rx x. a ie / 4
Fic. 1. Portulaca yecorensis, from the type collections 5 km NW of Yécora, Sonora, 29 Sep

1998, TR. Van Devender et al.. 98-1942, showing the large flowers and contrasting reddish
1mm.

foliage. Horizontal bar =

to the other, fleshy with membranous margins, obtuse at the cucullate tip,
slightly keeled above, but not crested below the tip, 3—3.5 mm long in flower,
enlarging to 5.5 mm long in fruit, the two sepals connate for 0.5 mm at
the base; petals 5, spreading, bright yellow, obovate, (4.5—)6-8(-9) mm
long, to (4-)5—6(—7) mm wide at the tip, strongly emarginate with an api-
cal notch to ca. 1.0—1.8 mm deep, the flowers averaging 14.7 mm in total
diameter (range 10-20 mm, n=30, field measured), the petals connate and
adnate to the sepals for about 0.4 mm at the sepal base; stamens (1 2—)15—
17; filaments 5.5—6.5 mm long, bright yellow, ascending, glabrous, adnate
to the petal-sepal bases for ca. 0.4 mm; anthers ca. 0.8—1.0 mm long, bright
yellow, with 4 elongate thecae, each ca. 0.2 mm in diameter (wet), the two
pair of anther sacs slightly twisted (1.e., not perfectly parallel); pollen glo-
bose, polyporate, (72—)90—98(—119) jim in diameter, light to strong yel-
low; style exserted beyond the stamens, bright yellow, 7.6—9 mm long, the
lobes 3-4, 1.5—2.5 mm long, the stigmatic surfaces long papillae with some
hairs to 0.3 mm long; ovules to 24. Ovary top obconic, to | mm long in
flower to 2.5 mm long, and 2.5 mm in basal diameter in fruit, the fruit
wall thin, with a slight constriction in the upper third; seeds (S—)10—19 per
ovary, reddish-black with a slight oil sheen, cucullate with a whitish patch

L004 Sipa 18(4)

Fic. 2. SEM of seed sculpturing patterns of Portulaca yecorensis: A. Edge view, showing series
of small papilli across the rim. B. Face view, showing outline of fruit, and the multiradiate
cells in the lower left. C. Oblique view, enlarged showing the cells with branched radii and
the caller papillate cells in the lower left. White bars = | mm in A and B, 0.3 mm in C.

where attached to the funiculus at the radicle bulge, 0.7—0.8(—0.86) mm
wide, to 0.5—0.56 mm thick, with some lateral cells tuberculate (Figs. 1, 2).
Additional specimens: MEXICO. Sonora. Municipio de Yécora: 2 km E of Yécora
28°22'SL"N, 108°54'53"W, 1600 m, 3 Sep 1996, A. Flores M. 4930, J. Sdnchez : (USON),
3.4km N of Yécora on road to Agua Blanca, oak woodland on bare volcanic hilltops, 28°29'35"N,
108°55'1L1"W, 1520 m, 23 Sep 1997, A. L. hae 97-1194, T. R. Van Devender, W. ars (ARIZ,
HUNT, NY, TEX, USON); ca. | km ESE of La Otra Banda (the Pima Indian eee of
42"N,

eee road to Talayotes, open pine-oak forest on locally bare mud flow surfaces, 21"
08°S54'1L2"W, 1600 m, 19 Nov 1997 A. L. Retna 97-1519, w - TR. Van See _M,
oe C. Cassa, A. E. Gondor (ARUZ): 2 km E of Yécora; 28°22'51"N, 108°54'53"W/, 1600

n, 3 Sep 1996, A. Flores M. 4930, J. Sdnchez (USON).

The new taxon appears related to the Portulaca oleracea complex. Like P.
oleracea, the plant is largely glabrous, moderately branched, and has crested
sepals, small axillary hairs, and yellow flowers. The new taxon differs from
P. oleracea in its much smaller size, smaller leaves, the distinctive reddish
color on all vegetative portions, the smaller and more sparse axillary hairs,
much larger flowers, larger pollen grains, poorly developed crest or keel on
the sepals, and seed shape and sculpturing (Fig. 1). The small size and smaller
leaves are, of course, a factor of its exposed, shallow-soil habitat and are fea-
tures not restricted to this species, as P. o/eracea, in similar habitats, may be
equally reduced. The red coloration caused by the presence of betalain pig-
ments in the outermost epidermis layer, is apparently a genetically fixed
feature allowing adaptation to the exposed, high-insolation habitat. Axil-
lary hairs are few and very short, measuring to 0.3 mm in length. In P. oleracea
they are slightly more conspicuous, extending to | mm in length. Porti-
laca oleracea has very small flowers (S—10 mm in diameter) while those of

_

—

HENRICKSON AND VAN DeveNpber, A new species of Portulaca 1005

the new species are 10-20 mm in diameter, as measured in the field by Van
Devender and Reina.

Portulaca oleracea is commonly reported as autogamous with flowers opening
for only about four hours during a day (Matthews et al. 1993; Geesink 1969).
The larger and more showy flowers of P. yecorensis are more conspicuous, a
feature that may be associated with attracting pollinators and possibly out-
breeding. The sphaeroidal, polyporate, pollen grains of P. yecorensis are very
large, measuring (72—)90—98(—119) jam in diameter; pollen of P. oleracea is
much smaller, 55—69 jm in diameter, but otherwise similar in structure.
Anthers in the new species are also much larger, 0.8—1 mm long (wetted).
Anthers of P. oleracea are about 0.5 mm in length (similarly wetted). Seeds
in the new species measure 0.68—0.83 in maximum diameter, which is comparable
to those of P. oleracea, but the body of the seed is more globose, not com-
pressed as in P. oleracea. The seed surface sculpturing pattern observed in
the new taxon gives evidence of its relationship. There are rows of cells on
the lateral surfaces of the body that are used to characterize the sculpturing
of the seeds (Danin 1978). The surface patterns found in the new species
are not entirely consistent, however, most lateral surface cells have a slight
metallic sheen and are stellate, to 0.2 mm wide, and have broad or narrow,
unbranched to forked or truncated radii that interlock with those of adja-
cent cells (Fig. 2). In some seeds this pattern is very similar to that of P.
retusa Engelm. (=P. oleracea L. subsp. impolita Danin & H.G. Baker) except
that distinct tubercules are absent except from some cells on the lateral walls.
In other seeds the sculpturing pattern is more obscure, but the pattern found
is more like that of P. retusa than those of P. oleracea. This leads me to con-
sider that the new species may be more closely related to P. retusa than P.
oleracea, if relationship indeed falls within this group. Matthews et al. (1993)
recently combined P. retusa with P. oleracea; the senior author is in strong
disagreement with this action.

The contrasting dark red foliage and large, bright yellow petals make
the plant very conspicuous. Red vegetative pigmentation is a common stress
response in many succulent and non-succulent species (e.g., Amaranthus,
Portulaca, Sedum) in answer to high insolation. This commonly develops over
a season as light stress increases. However, its development can be controlled
genetically. Danin et al. (1978) note that some cultivated subspecies of P.
oleracea characteristically developed pigmentation of the sepals, etc., while
other subspecies would not, which would indicate genetic control of pig-
ment placement. The red pigments are betalains (Clement & Mabry 1996)
and, in this taxon, are largely confined to epidermal layers. Such red pig-
ments would reflect red and absorb green wavelengths of light, while the
green chlorophyll pigment would absorb the remaining red and reflect green

1006 Sipa 18(4)
wavelengths; this to some extent reduces absorptivity of solar radiation (Von
Willerc et al. 1992). In areas of high insolation, however, this does not impede
the light available for photosynthesis. The result is dark (apple) red or maroon
foliage. In the new species all vegetative portions of the plant (stems, leaves,
sepals) had the distinctive, dark red coloration and this developed from the
beginning—it was present throughout seedling stage into the adult plane,
irregardless of the amount of light given to the plant. Even seedlings grown
in the senior author's shaded, north-facing office window developed and retained
the characteristic apple-red pigmentation on all vegetative structures.

The mudflow barren habitat is so open that the plants are fully exposed
throughout their two to three month life span. Surveys in this habitat on
August 17, 1998, revealed seedling Sedum vinicolor but not Portulaca yecorensis.
Peak Howering collections were made on September 23, 1997, and September
29, 1998, nearing the end of the summer monsoon rains. The November
19, 1997, collection was mostly of dead, dried plants. It is interesting to
note that the life cycle of P. yecorensis appears to be delayed compared to the
annual composites (Pectis vandevenderi, Tridax yecorana) which senesce in October.
Likely the succulent leaves allow it to persist longer than non-succulent
annuals after che summer rains taper off.

on

ACKNOWLEDGMENTS
Neil Harriman provided the Latin translation. Richard Felger, Robert Kiger,
Phil Jenkins, and Myron Kimnach gave helpful early suggestions as to the
plant’s identity. Agustin Flores M. and Jestis Sanchez E. of the Universidad
de Sonora collected the plants in 1996 and stimulated our interest in it.
Father William Trauba of Yécora helped in all aspects of the field work. Bob
Scarborough provided information on the geology of the mudflow surfaces.

REFERENCES

Burqurz, M., A.A. Martinez Y. and P.S. Martin. 1992. From the high Sierra Madre to
the coast: changes in vegetation is highway 16, Maycoba-Hermosillo. In: K.P. Clark,
J. Roldan, and R. H. Schmidt, eds. Ge ‘ology and mineral resources of the northern Si-
erra Madre Occidental. Guidebook, El Paso Geological Survey Publication No. 24, El
Paso. Pp 239-252.

CLEMENT, J.S. and T.J. Masry. 1966. ee evolution in the Caryophyllales: a system-
wcic overview. Bot. Acta 109:360

CocHeme, J.and A. Demanr. 1991. i g ‘the Yécora area, northern Sierra Madre Occidental,
Mexico. Geol. Soc. America Special Pap. 2

Danin, A., 1. BAKER and H.G. - AKER. 1978. Cae and taxonomy of the Portulaca
oleracea L. complex. Israel J. Bot. 27:177—211.

GEESINK, R. 1969. An account - the genus Portulaca in Indo-Australia and the Pacifi
(Portulacaceae). Blu L727 5=301.

Martruews, J.F., DLW. Ken NTRON, and $.F. ZANE, 1993. The biology and taxonomy of Portu-
laca oleracea L. (Portulaceace) complex in North America. Rhodora 95:166-183

HENRICKSON AND VAN DeveENDER, A new species of Portulaca 1007

Reina, G.A.L., T.R. VAN DevENDeER, W. Trauba, and A. BuUrQquEz M. 1999. Caminos de
Yécora. Notes on the vegetation and flora of Yécora, Sonora. In: M. Ortega N., ed. Sym-
posium internacional sobre la utilizacién y aprovechamiento de la flora de zonas aridas,
Universidad de Sonora, Hermosillo.

Von Witert, D.J., B.M. Etter, M.J.A. WERGER, E. BRINCKMANN, and H.D. IHLENFELDT.
1992. Life strategies of succulents in deserts. Cambridge University Press, Cambridge.

L008 Sipa 18(4

—

BOOK REVIEW
RopBert H. MOHLENBROCK 1999. The Illustrated Flora of [linois: Ferns.
Second Edition. (ISBN 0-8093-2255-2, hbk.). Southern Illinois University
Press, P.O. Box 3697, Carbondale, IL 62902-3697. Tel.: 618-453-6633;
Fax: 618-453-1221; email: danseit@siu.edu. $39.95. 240 pp., 163 illus.

This second edition of a truly classic floristic work includes 25 species not previously re-
ported in che 1967 edition, has additional illustrations for all of those taxa, and provides a
key to the additions and to supplement the other keys in the main section of the book
when (as in the Eysepeds) Benen concepts in the family have changed. The small size of
the book and its simple terminology, concise descriptions, literature citation, Common name,

habitat, range and distribution within the state, all make this an ideal field tool, just as its
author intended. The simple yet clear illustrations, with accompanying dot-distribution
maps are also very useful.

Phe introduction and keys to the orders and also, the genera, of ferns and their allies
contain information requisite for any pteridology course. The Appendix of 47 pages, in-
cluding additions, changes to keys, descriptions of taxa not previously included, distribu-
tion maps, etc. 1s a little cumbersome, but I can imagine that ic would have been extraor-
dinarily time-consuming to have integrated the updated information into the appropriate
niche within the main text body. The glossary is short, simple and concise.

Certainly, the first edition of this book is already dog-eared, taped many times with li-
brary repair tape, and otherwise patched, on the shelves of everyone who botanizes in the
vicinity of Illinois. Certainly I will have mine in the car when I go back to those septentrional
stomping grounds tn and around the Land of Lincoln. There is nothing as welcome as an
update to a classic and this book fits the bill perfectly. I am delighted chat Mohlenbrock
was able to update this classic work so that younger botanists and “Phern Physicians” might
better enjoy the marvelous fern flora. Southern Illinois Press deserves a salute for printing
this fine book; truly a bargain!!! I hope

more institutions in the state and adjacent ones
will consider teaching a pteridology course if they are not presently doing so, given the
importance of the group and its abundance in the state.—-John J. Pipoly IL

Stipa L8(4): 1008. 1999

REVIEW OF EARLY NOMENCLATURE IN
EUTHAMIA (ASTERACEAE: ASTEREAE)

GUY L. NESOM
Biota of North America Program
North Carolina Botanical Garden
Coker Hall CB 3280
University of North Carolina
Chapel Hill, NC 27599, U.S.A

ABSTRACT

A review of the early nomenclature of Exthamia s riz bservations by Sieren (1981)
and Reveal (1991) and adds other perspectives. Nuttall published the group in 1818 in
ee format of a new subdivision of see interpreted here at sectional rank, but noted it
as “a subgenus, or rather genus.” Cassini in 1825 provided the elements for validation of
ls at generic rank, ee he did so pee ee and ascribed the name to Nuttall.
In this interpretation, separate citations of lectotype and type are required for the two names
(sect. Sok Nutt. and genus Evthamia Nutt. ex Cass., respectively). Also in this inter-
pretation, Nuttall in 1841 validated nomenclatural combinations for E. SrgTeT ACI 7 )
Nutt. and E. tenuifolia (Pursh) Nutt. and added E. occidentalis Nutt. as a new species

me Exthamia caroliniana (L.) Greene ex Porter & Britt. was validated in 1894, — on
Er rigeron carolinianus L., and includes E. tenuifolia as a synonym. There 1s reason to maintain
Euthamia galetorum Greene at specific rank rather than as a variety of E. caroliniana or E.
graminifolia. The discussion includes application of several sien controversial inter-
the term type or an equiva-

«6

pretations of the ICBN: disposition of “alternative names;
lent” in typification; - conditions of “indirect reference” for va lid Seed.

RESUMEN

Una revision de la nomenclatura previa de Evthamia resume las observaciones de Sieren
(1981) y Reveal (1991) y afiade otras perspectivas. Nuttall publico el grupo en 1818 como
una nueva Supa isin Salida, que se interpreta aqui en el rango de secci6n, pero la cit6

a éner z4s género.” Cassini en 1825 dio los elementos para la validacion

como “un
de fallen en a rango eno. pero lo hizo asi sin querer y adscribié el nombre a Nuttall.
En esta se requieren le lectotipo y tipo para los dos nombres

(sect. Buamic Nutt. y género Evthamia Nutt. ex Cass., respectivamente). También en esta
interpretaci6n, Nuttall en 1841 valid6 combinaciones nomenclaturales cone graminifolia
(L.) Nutc. y £. fenafoua Bussh) Nutt. y afiadié E. ocerdentalis Nutt. como una nueva especie.
El nombre Euthamia caroliniana (L.) Greene ex Porter & Britt. fue validado en Geos. basado

n Erigeron carolinianus L., e incluye E. tennifolia como sinénimo. Hay una raz6n para mantener
Eilune ae Greene en el rango especifico en vez de variedad de E. caroliniana o E.
eraminifolia. La discusi6n incluye la aplicaci6n de varias interpretaciones potencialm
ee del ICBN: disposici6n de “nombres alternativos;” “el término tipo o un ee
en la tipificacién y condiciones de “referencia indirecta” para la publicacién valida.

Reveal (1991) clarified the application of the Linnaean name Erigeron

Sipa 18(4): 1009-1018. 1999

LOLO Sipa 18(4)

carolinianus L., confirming earlier opinions that it represents a species of
Euthamia and that this species is correctly treated as Euthamia caroliniana
(L.) Greene ex Porter & Britton, of which Evthamia tenuifolia (Pursh) Nutt.
is asynonym. Reveal also noted that the name for Evthamia at generic rank
should be cited as Evthamia (Nutt.) Nutt. ex Cass. (see below) and that the
choice of lectotype for the genus by Britton and Brown in 1913 should stand
as Exthamia graminifolia (L.) Nutt. Except for details regarding the generic
authorship and typification and the authorship of E. tenwifolia, his observa-
tions are firm, but several other associated nomenclatural points were left
unresolved, these discussed here. Previous perspectives regarding author-
ship are summarized, additional comments are provided, and a nomencla-
tural summary of the names involved follows at the end of the comments.

Other published comments regarding Evthamia on a more limited or re-
gional basis have not touched upon the nomenclatural topics discussed here.

Euthamia at generic rank

Euthamia was ambiguously described by Nuttall (1818, p. 162) as “A sub-
genus, or rather genus, reciprocally allied to So/idago and Chrysoma.” Vari-
ous botanists (e.g., Elliott 1823; de Candolle 1836; Greene 1902; Gleason
& Cronquist 1991, Cronquist 1994, as inferred from citation of the genus
simply as “Exthamia Nutt.”) have tacitly accepted 1818 as the validation
date for the genus, but others have not (e.g., Nuttall himself in 1841; Sieren
1981, monographer of the genus, who cited “(Nutt.) Nutt.” as the author-
ity, regarding 1841 as the date of validation). The only other previous, ex-
plicit, and published consideration (Reveal 1991) of this problem concluded
that the elements of validation were not provided in 1818. The implica-
tion of the 1994 ICBN Article 34.2 regarding identical, simultaneously
published names at different ranks (“alternative names”) after | January 1953
appears to be that such names are not necessarily invalid if published be-
fore that date. The position here, however, and that perhaps implicitly taken
by Reveal, is that a name at only one of the ranks should be recognized as
valid, if a balance of evidence regarding the author's intention suggests a
resolution to the ambiguity.

Nuttall in 1818 placed two species under the heading of Euthamia, which
was marked by an “asterisk” (indicating a new taxon) near, but before, the
end of the So/dago treatment. Thus Evthamia was not in the numbered se-
quence of genera treated by Nuttall (So/idago, including Exthamia, is genus
560; Brachyris is genus 561). Nor is Exthamia included in the index to ge-
neric names in the Genera of North American Plants. The two Euthamia
species are numbered “50” and “51,” terminating the numbering sequence
for species of Solidago. After the comments on Exthamia, Nuttall returned
to Solidago in a summary paragraph (provided ina manner and position similar

Nesom, Review of early nomenclature in Euthamia 1011

to such summaries for many other genera) that noted the overall geographic
distribution of the genus and suggested that arborescent species of St. Hel-
ena and New Zealand probably should be excluded from Solzdago. In con-
trast to the other species of So/idago, however, and lending to the ambigu-
ity, Nuttall provided basionyms and author and publication citations for
the Evthamia species, listing “50. graminifolia. Chrysocoma graminifolia” and
“51. cenuifolia. S{olidago}. tenuifolia,” giving an impression of his intention
to make new nomenclatural combinations for these two species. The man-
ner in which the treatment of Evthamia is imbedded within So/zdago, how-
ever, is viewed here as more explicit evidence for regarding Evthamia of 1818
at infrageneric rank.

Nuttall’s description of Evthamia as a “subgenus, or rather genus” was in
the second paragraph of description following the “EUTHAMIA” heading,
but the format for the delimitation of Ezthamia is identical with other infrageneric
names proposed in the same volume and long-accepted at sectional rank
(see comments below, “Evthamia at infrageneric rank”). Nuttall (p. 151)
applied almost exactly the same description to his group “II” of Inu/a sect.
Chrysopsis, noting that “This genus, or subgenus, appears to be peculiar to
North America.” This heterogeneous group (as now seen) was composed of
white-rayed species of Aster (compared to typical yellow-rayed Chrysopsis)
with a double pappus, and it seems clear that the description here of “ge-
nus, or subgenus” was meant to be taken informally. The situation in Euthamia
is analogous: the species were treated as a section within Solidago, but Nuttall’s
accompanying comment suggests that a higher rank for them is reasonable.
In fact, it is the directness of this suggestion (and its close proximity to the
listing of the new name) that has created the ambiguity of interpretation.

More than 20 years later, Nuttall (1841, pp. 325-326) provided a more
definite account of Evthamia, explicitly treating it at generic rank. Here he
cited the basionym for Exthamia “Asa section of Solidago, Nutt., Gen. Am.,
Vol. I., p. 162. Decand. Prod., Vol. V., p. 341.),” indicating that he re-
garded the Evthamia of his 1818 publication to have been at infrageneric
rank, as was Euthamia of de Candolle. Nuttall apparently intended to rec-
ognize the name at generic rank, with the authority understood to be “(Nutt.)
Nutt.” Remarkably, however, his ambiguity regarding the status of the name
was perpetuated even here, as he did not provide Euthamia (in the “header”)
with an asterisk characteristic of the new names (e.g., Ericameria Nutt.) and
new combinations (e.g., Amphiachyris (DC.) Nutt.) at generic rank elsewhere
in the treatise. His treatment of E. graminifolia and E. tenuifolia (see below)
also might be taken to imply that he assumed those names had already been
incorporated into the valid nomenclature for Evthamia.

Meanwhile, as pointed out by Reveal, Cassini (1825) had preceded Nuttall’s
1841 comments in providing the elements of validation for Evthamia at

LO12 Stipa 18(4)

generic rank. Here Cassini referred to “Le genre Ezthamia de M. Nuttall” (a
clear but indirect reference), apparently assuming that Nuttall in 1818 had
effectively established it at generic rank and giving Nuttall credit for its
publication. In his associated description and comments, Cassini unambiguously
treated Exthamia as a genus, although he surely did so without the speci

—
a)

ic
intention of publishing a formal validation at that rank. In contrast to the
suggestion of Reveal (1991) that the authority for the genus Evthamia be
cited as “(Nutt.) Nutt. ex Cass.,” its citation simply as “Nutt. ex Cass.”
acknowledges Cassini’s role in validating the name as well as his explicit
recognition that the name should be ascribed to Nuttall. This appears to
be in accord with guidelines for citation outlined in Article 46.4 of the 1994
ICBN. Cassini did not furnish the necessities for any nomenclatural com-
binations in Evthamia at specific rank.

—
on

In summary, evidence indicates the name Euthamia was ae pub-
ned at infrageneric rank (interpreted here to be sectional), and despite
Nuttall’s ambiguity both in 1818 and 1841, the most explicit parts of his
treatments suggest that the validation of Bahan at generic rank was ef-
fected by Cassini (as “Nutt. ex Cass.”). Or, if Cassini's procologue were con-
sidered an insufficient basis for validation, Sieren’s interpretation could be
followed by citing the authority as “(Nutt.) Nutt.” Ocher interpretations
would read the evidence as favoring validation of the name in 1818 as a
genus rather than section or subgenus or else perhaps find the evidence so
evenly equivocal that the name from 1818, with Nuttall as sole author, could
be regarded as valid at two or even three ranks (as noted above, according
to ICBN Article 34.2). Or, perhaps any degree of ambiguity should lead to
the formal acceptance of alternative names.

—_

lis

Euthamia at infrageneric rank
Although Nuttall’s descriptive phrase for Evthamia in the 1818 publica-
tion was “a subgenus, or rather genus,” he stated in 1841 that Euthamia
was positioned “as a section of So/idago” in 1818. The latter rank is accepted
here for the original publication of the name, as it is Nuttall’s most unam-
biguous taxonomic characterization and one that is the most consistent with
his 1818 format for analogous names in other genera. Other supraspecific
taxa, securely accepted at sectional rank, were described by Nuttall in 1818
with exactly the same format as Euthamia (e.g., Inula sect. Chrysopsis Nutt.,
p. 150; Erzgeron sect. Caenotus Nutt., p. 148; see Semple 1981 and Cronquist

1947, respective ly). Each of these names was given in small-sized capital
letters, preceded by an asterisk to indicate that it was a newly proposed name,
and followed by a period and a dagger, referring to a footnote providing the
name’s derivation.

Alternatively, in view of ambiguity regarding the rank of Nuttall’s 1818

Nesom, Review of early nomenclature in Euthamia 1013

Euthamia, it nevertheless may be considered validly published as an unranked
subdivision of So/zdago, according to ICBN Article 35.2: “A new name or
combination published before 1 January 1953 without a clear indication of
rank is validly published provided that all other requirements for valid publication
are fulfilled; ... it may serve as a basionym or replaced synonym for subse-
quent combinations. ...

De Candolle (1836, p. 341) explicitly treated Euthamia as a section of
Solidago, citing both Nuttall (1818) and Cassini (1825) as having treated it
at generic rank, although he acknowledged that the group might be ac-
ceptable as a genus (“An genus proprium ut innuit Nuttall et asserit Cassini?”).
De Candolle’s choice of a less than fully forceful verb (“innuo”) for Nuttall’s
description appears to signal a degree of uncertainty about the rank, espe-
cially as the phrase is ended with a question mark.

Erigeron sect. Multiflori G. Don in Loudon (1830) comprised only two
species in its original description, Evrigeron villarsii Bell. (= Erigeron atticus
Villars) and Erzgeron carolinianus (= Euthamia caroliniana). Sect. Multiflora

was lectotypified (Nesom 1989) by Erigeron carolinianus partly to avoid dis-
placing the widely used Evigeron sect. Trimorpha (Cass.) DC. (Prodr. 5:290.
1836), of which E. villarsii is a member, and partly because, at the time,

the identity of Erigeron carolinianus appeared to have little chance of being
removed from the realm of ambiguity.

Discussion of infrageneric categories within Euthamia is largely academic,
because the distinctiveness of the genus, apart from any other, is now gen-
erally accepted, and marked homogeneity among the relatively few species
(6-8 total) suggests that formally designated categories will hardly be nec-
essary or useful.

Typification of Exthamia
If valid publication of Euthamia at generic rank is attributed to Cassini as a
new name (as in the interpretation here) rather than a new combination,
typification of the genus also was effected by Cassini. His technical description
of Euthamia (1825, p. 471) was explicitly drawn from Chrysocoma graminifolia
L. (en traccant ici les caractéres genériques observés par nous sur la Chrysocoma
gramifolia de Linné”) and only that species. The position of this species in
fixing the application of the name Euthamia in Cassini’s discussion is un-
ambiguous. At the time of Cassini’s work, his short phrase “indicating the
generic characters,” in reference to C. graminifolia, was a clear and concise
“equivalent” to the term “type,” as that term is understood today in bo-
tanical nomenclature, apparently satisfying the requirements of the 1994
ICBN (Article 7,11),

In the description of Solidago sect. Euthamia, Nuttall in 1818 did not
specify which of the two included species should serve as the type. Nor did

L014 Sipa 18(4)

he in 1841 make a choice among the three species included in Evthamia at
generic rank. De Candolle (1836) included only two species in his treat-
ment of Solidago sect. Euthamia and did not indicate which was to be re-
garded as the type. As noted by Reveal, Britton and Brown (1913) cited E.
graminifolia as the generitype, which can be taken as an effective lectotypification;
Sieren (1981) cited EF. tenuifolia as the generitype. Alternately, if Reveal’s
interpretation of the validation of Evthamia as a new combination by Cassini

were accepted, Cassini's presentation and documentation apparently can be
be taken as the first effective lectotypification, based on Chrysocoma graminifolia
(= Euthamia graminifolia), assuming that he provided an acceptable equiva-
lent to the term “type.

Discussions of the process of lectotypification and its formal codal (ICBN)
requirements have outlined ambiguities of interpretation (e.g., Barrie et
al. 1992a, 1992b; Winter et al. 1992; Zijlstra 1992). A summary example
of the problem of “the term type or an equivalent” is given in Brummit
(1994), referring to a proposal by Reveal (1991). The course suggested here
for the lectotypification of “Exthamia (Nutt.) Cass.” does not appear to be
contradicted by the current Code (ICBN 1994), unless unwritten interpre-
tations or implications are brought to the fore. In any case, Evthamia graminifolia
is the lectotype of any name based on ea sect. Euthamia Nutt., whether
designated by Cassini or by Britton & Brow

Authorship of Evthamia species

Various botanists have used the names Evthamia graminifolia (L.) Nutt. and
Euthamia tenuifolia (Pursh) Nutt., interpreting Nuttall’s epithets from 1818
as validly published in Exthamia. In one of the earliest examples, Elliott
(1824) cited both names as such in lists of synonyms under their accepted
names in So/idago. Greene, both in 1894 (in Porter and Britton) and in his
later overview of the genus (1902), regarded 1818 as the date of valid pub-
lication for the genus and for Nuttall’s two names at specific rank. Sieren
(1 98 | ) regarded E, graminifolia as validated by Nuttall in 1841 but E. tenutfolia
in 1818. Cronquist (1980) referred to E. tenuifolia (Pursh) Greene but later
(in Gleason and Cronquist 1991) changed the citation to E. renwifolia (Pursh)
Nutt. Reveal (1991) regarded the latter to have been validated by Greene
(1902) as E. tennifolia (Pursh) Greene.

In Nuttall’s 1841 treatment of Evthamia, he included three species, E.
graminifolia, E. tenuifolia, and E. occidentalis (sp. nov.), each epithet associ-
ated with the generic name. Nuttall did not cite basionyms or citations of
earlier publication for FE. graminifolia and E. tenuifolia, but it is clear that he
was referring to the taxa originally published by Linnaeus and Pursh, re-
spectively, as an update of the descriptions in his 1818 publication. The

1841 descriptions for these two species are rewritten and somewhat expanded

Nesom, Review of early nomenclature in Euthamia 1015

compared to the earlier ones, where the earlier sources of the names were
cited. In contrast, E. occidentalis Nutt. was marked as a newly proposed name
by an asterisk preceding the epithet, this symbol lacking from the other
two names. Despite omission of basionyms for the first two species, Nuttall’s
1841 treatment of Exthamia provided a specific reference to his 1818 pub-
lication, and there can be little doubt that he regarded the first two names
in Exthamia to be in parallel with those of his previous manuscript, which
included the basionyms and publication citations. This appears to satisfy
the condition of “indirect reference” for valid publication of names (1994
ICBN Articles 32.4 and 32.5), and the validation of E. graminifolia and E.
tenuifolia in 1841 is accepted here.

After Nuttall’s formal treatment of Evthamia in 1841, this group of plants
apparently was not again recognized at generic rank until 1894, when Por-
ter and Britton (1894) provided E.L. Greene’s view of the group in a formal
nomenclatural summary for the species in northeastern North America. The
three species listed were E. caroliniana (including E. tenuifolia as a synonym),
E. graminifolia, and E. leptocephala; the first and third are regarded here as
receiving formal validation at specific rank in the 1894 publication. In a
more comprehensive treatment of Exthamia, Greene (1902) separated the
concepts of Euthamia caroliniana and E. tenuifolia, but it is now generally
acknowledged that the types of these two names represent a single species.

Status of Euthamia galetorum
With acknowledgment that the correct name of Exthamia tenuifolia is E,
cavoliniana, a decision is required regarding a varietal combination within
E. tenuifolia. Euthamia galetorum Greene has been treated as a variety of both
E. tenuifolia (Fernald 1921, as Solidago tenuifolia var. pycnocephala Fern.) and
E. graminifolia (House 1924, as Solidago graminifolia var. galetorum (Greene)
House). Friesner (1933) and Harris (1943), as well as the recent monographer
of Exthamia (Sieren 1981), maintained E. galetorum at specific rank. While
Roland and Smith (1969) noted that S. tenuifolia and S. galetorum are “evi-
dently closely related,” they also maintained both entities at specific rank.
In contrast, Taylor and Taylor (1983) formalized the varietal status of E.
galetorum within E. tenuifolia (as var. pycnocephala (Fern.) C.&J. Taylor), not-
ing that “field studies along with examination of types and other herbarium
specimens support Fernald’s treatment as a variety of tenuifolia” (p. 178).
Fernald (1921, pp. 143-144) observed that Solidago tennifolia vat. pycnocephala
is “a very distinct goldenrod ... everywhere dominant [in southern Nova
Scotia} and thoroughly characteristic of these sandy and cobbly lake-margins”
but that “Too many collections ... show direct transition [to var. tenwifolia] in
all these characters to allow the specific separation of the Nova Scotian plant.”
Most of the intermediacy described by Fernald involves features of habit

LO1G6 Stipa 18(A4)

and leaf morphology. In later descriptions, Fernald (1950) noted that heads
of E. galetorum have 25—50 flowers, in contrast to the 12—20 flowers per
head in E. tenuifolia (this comparison modified to 20—50 vs. 10—20 by Sieren).

Sieren (1981, p. 560) noted that Exthamia galetorum “is readily separated
from {E. tenuifolia} by its large numbers of flowers, especially the disc, its
wider, ascending leaves, and the absence of axillary fascicles. In its gross
morphology, E. ga/etorum most closely approaches the wide, bluntish-leaved
variety of E. graminifolia, variety major.” In specimens of E. galetorum I have
examined, its few-branched stems, relatively short leaves, and small, com-
pact inflorescences of few, large heads with numerous flowers, appear to be
distinct from E. teniifolia in the same region, in agreement with the view
of Sieren and others. Until stronger evidence is presented to counter the
observations and broadly based consideration of Sieren, it seems reasonable
to maintain E. ga/etorum at specific rank rather than placing it varietally

under E. caroliniana.

Euthamia galetorum was known only from Nova Scotia by Sieren (1981),
but it has subsequently been identified from Maine and New Hampshire
(Bruce Sorrie, pers. comm.), and House (1924) described the entity from
various parts of New York, distinguishing it from E. fenwifolia and other
related taxa.

NOMENCLATURAL SUMMARY

Taxa listed are those included in the present discussion. Those in bold,
with accompanying authorship, are as accepted in the interpretation here.
Euthamia Nutt. ex Cass. in Cuvier, Dict. Sci. Nat. 37:471. 1825. Type: Chrysocoma

graminifolia L. (= Euthamia graminifolia (L.) Nutt.).

Solidago sect. Euthamia Nutt., Gen. N. Amer. PI. 2:162. tStS. Lectotype: Exthamia graminifolia

(L.) Nutt., as designated by Britton & Brown (191

Enthamia (Nutt.) Nutt., Trans. Amer. Philos. a ser. 2, 7:325. 1841. (nom. superfl.).

Solidago sect. pera .) DC., Prodr. 5:34 et Gone superfl.).

Solidago subg. Exthamia (Nutt.) House, Bull. wa York State Mus. 254:694, 1924.

Erigeron sect. Submultiflori G. Don in Loudon, Hort. Brit. 343. 1830. Lectotype: Erigeron

carolinianus L. (= Euthamta cavoliniana (L.) Greene ex Porter & Britton), as designated
by Nesom (1989).

Euthamia caroliniana (L.) Greene ex Porter & Britton, Mem. Torrey Bot.
1894

Club. 5421,
Evigeron carolinianus L., Sp. ‘ 863
Solidago caroliniana (L.) B.S 2 Ril an C at. N.Y. 26. 1888.
Solidago tennifolia Pursh, F ‘ er. Septent. 2:540. i: 14

Euthamia tenunifolia ( (Purch) cue Trans. Amer. Philos. Soc. ser. 2, 7:326. 1841

Euthamia galetorum Greene, Leafl. Bot. Observ. Crit. 2:152. 1911.

Solidago tennifolia var. pycnocephala Fern., Rhodora 23:293. 1921.

Nesom, Review of early nomenclature in Euthamia 1017

Enthamia tenuifolia vat. ee (Fern.) C. & J. Taylor, Sida 10:177. 1983.
Solidago graminifolia var. galetorum (Green e) House, N.Y. State LS Bull. 243—244:45. 1923.
Solidago galetorum (eee ae fades Univ. Bot. Stud. 3:58. 1933.

Euthamia graminifolia (L.) Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:325.
1841

Chrysocoma graminifolia L., Sp. Pl. 841. 1753.
Solidago graminifolia (L.) Salish. Prodr. 109. 1796.

y. Bot, Club 5.321.
Solidago a Torr. & Gray, Fl. N. Amer. 2:226. 1841.

suas leptocephala ae & Gray) Greene ex Porter & Britton, Mem.
T 894.

Euthamia occidentalis Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:326. 1841.

Postscript.—After review and revision of the present manuscript, a com-
mentary by K.N. Gandhi appeared in print, covering many of the same topics
and reviewing similar rationale. Gandhi (1999) also concludes that the 1818
publication of Exthamia was at infrageneric (but subgeneric) rank and credits
Cassini with its validation at generic rank, although he interprets the au-
thorship as “(Nutt.) Cass.,” noting that Cassini's indirect reference to Nuttall’s
earlier work brings the basionym into consideration. As noted by Gandhi,
this brings the number of possibilities for formal citation of the generic
authorship to four. Validation of E. graminifolia and E. tenuifolia is attrib-
uted to Nuttall in 1841, as in the interpretation here.

,

ACKNOWLEDGMENTS
Iam grateful to Rogers McVaugh for his ideas regarding various points
discussed in the present paper, Jim Reveal for information on concepts of
lectotypification, Ken Wurdack for comments on an early version of the
manuscript, and to an anonymous reviewer for a different point of view,
which helped to focus the presentation here. A discussion with John Strother
also illuminated different interpretations and was he pfu

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A NEW BULBOPHYLLUM (ORCHIDACEAE)
SPECIES FROM THE CRATER MOUNTAIN
AREA IN PAPUA NEW GUINEA

N.H.S.HOWCROFT

P.O. Box 30, Keravat
East New Britain, PAPUA NEW GUINEA
howcroft@daltron.com. pg

ABSTRACT

A new species Bulbophyllum wakoi, section Brachystele Schltr., collected from the Crater
Mountain Wild Life Management Area in Papua New Guinea is described. It resembles B.
foetidum Schltr. and its variety B. foetidum Schltr. var. grandiflorum J.J. Smith. However, it
differs from these in the size and the shape of the sepals, petals, labellum, and column arms.

ABSTRACT (MELANESIAN TOK PISIN)

Niupela kain orchid Bulbophyllum wakoi, sekson Brachystele Schltr., ol bin kisim long
Creta Mauntein Wail Laif Management Eria insait long Papua New Grinaee m ol bin diskraibim.
Em i wankain long B. foetidim Schltr. na wantaim variety B. foetidum var. grandiflorum J.J Smith.
Dispela em minim olsem long dispela insait long sais na seip long sepals, petals, labellum,
na colum ams.

INTRODUCTION

The section Brachystele Schltr. in New Guinea comprises about three taxa.
These are B. foetadum Schltr., B. foetidum Schltr. var. grandiflorum J.J.Sm. and
B. pachyanthum Schltr. The Crater Mountain species is closely related to B.
foetidum Schlitr., to which it has been compared with, but differs in size and
shape of the floral parts and more specifically in the some features of the
column and the labellum. The newly recognised taxon is described here.
Bulbophyllum ac! “leas sp. nov. (Fig. 1). Tyeus: PAPUA NEW GUINEA.

Simbu Province: Ex Herowana, Crater Mountain Wildlife Management Area, S. Wako
105, 10 Apr ne en s: LAE 79039).

Species nova affinis B. foetidum Sc nea et - ee var. nea J.Smith sed floribus
majoribus, sepalis et petalis latioribus, | g et majoribus, brachiis

spathulatis et majoribus.

Epiphytic, the creeping and branching habit typical of the section. Psewdobulbs
ca. 34cmx 1.4cm, oblong, transversely elliptic, slightly flattened. Leaves petio-
late, the petiole semi-conduplicate. Inflorescence 33 cm or more long, 9 or
more flowers; rachis short, 2.5 cm long, glabrous, nodding; peduncle more
than 31 cm long, 0.59 cm in diameter, sheaths more than 4, glabrous, tubular;
floral bracts ovate to oblong-ovate, 2.7 cm long, 1.2 cm wide, concave, acute,
glabrous. F/owers positioned in all directions, more or less patent, very large,

Sipa 18(4): 1019-1023. 1999

1020 Stipa L8(4)

Fic. |. Bulbophyllum wakoi. A. Leat and apex of pseudobulb. B. Inflorescence, 5 cm scale. C.
Flower. D—G. Floral bract, dorsal sepal, lateral sepal, petal, | cm scale. H. Column and
labellum, lateral view. I-K. Labellum back, dorsal and ventral views, | cm scale. L—M.
Column, ventral and lateral view.

Howcrort, A new Bulbophyllum species in Papua New Guinea 1021

moderately to wide opening; dorsal sepal ovate, 4.4 cm long, 1.6 cm wide,
glabrous, tip acuminate, semi-conduplicate slightly recurved; lateral sepals
free, obliquely triangular 5.0 cm long, 2.1 cm wide; glabrous, lower basal
margin obliquely triangular or lobed, tip acuminate, se oneu aca, petals
ovate-lanceolate to elliptic, slightly concave, glabrous, 3.7 cm long, 1.1 cm

wide, tip long acuminate; /abellum mobile, thick, fleshy, in general outline
cordiform or broadly ovate, 1.8 cm long, 1.2 cm wide near base, apex blunt
but ventrally bilobed, convex above with a longitudinal groove producing
2 slight longitudinal ridge and terminating ina mildly raised cushion at apex,
margin slightly papillose; adaxial side slightly concave with an inconspicuous
median ridge; column dorsally slightly curved, from ovary to tip of stelid
1.2 cm long, to tip of rostellum 1.0 cm long; dorsal stelid short and blunt,
stelids on lower margin conspicuous, subulate; basal arms of column por-
rect, equal in length to stelids, spathulate, from column foot ca. 1.0 cm
long, ca. 0.5 cm broad; column foot at the right angles to column, ca. 1.0 cm
long curved towards base, broad, quadragular, truncate ca.1.0 cm wide, with
a lateral falcate lobe, internal face longitudinally convexed with a raised
ridge; anther hooded; stigma more or less quadrangular; ovary ca. 1.0 cm long,
glabrous; pedicel ca. 4.0 cm long, glabrous; pol//en not seen; fruit not seen.

Distribution.—Papua New Guinea. Herowana area in the Crater Moun-
tain wild life management area, Simbu Province.

Habitat.—Upper montane rainforest zone. No records accompanied the
holotype specimen, but all the New Guinea species come from areas with
rainfalls between 2,000-4,000 mm per annum. Species of this section Brachystele
are found growing in the thick root and leaf litter of rainforests and as epi-
phytes climbing the trunks of trees. Information on altitude was not avail-
able but Herowana is located at 850 m altitude and Crater Mountain rises
to 2100m. The most likely altitude for this species is around 1,500 m.

This new species is a closely allied to B. foetidum Schltr. (Fig. 2) includ-
ing B. foetidum var. grandiflorum J.J.Sm., but has larger lowers, wider sepals
and petals; the labellum is not oblong but is cordate and larger; arms of
column are spathulate and larger. The most important differences between
B. wakoi and B. foetidum lie in the shape of the labellum (Figs. 1 C, I, J and
Figs. 2 B, H—J) and the differences in the size, shape and posture of the
basal arms of the column (Figs. 1 H, Land Figs. 2 FR M,N, L; Figs. 2 M and
L represent two specimens from different locations).

The illustration of B. foetidum by Schlechter (1913) and Smith’s illustration
of variety grandiflorum (1929) compare fairly well with Figure 2, but the
differences between the basal arms of the column, in length and shape, in both
illustrations, do not fit that of B. wakoi. Smith's variety was also described
as having pustules on the rachis. These are not present in the new species.

Bulbophyllum foetidum, as the name suggests, produces as foul odor. When

1022

Sipa 18(4)

: edt

Fig. 2. Bulbophyllum foetidum. A. Habit, 5 cm scale. B. Flower. C—E. Lateral sepal, dorsal
sepal, petal, 1 cm scale. F. Column and labellum, lateral view. G-H. Labellum, lateral and

orsal views. I—J. Labellum, back view of two speciemns. L—-M. Column, ventral view
ol post specimens. N. Column, 1 cm scale. O. Anther cap, lateral
&

=

Oomsis and Morobe patr
and front views (lefe to right

Howcrorr, A new Bulbophyllum species in Papua New Guinea 1023

extracting the specimen of B. wekoi from the alcohol, in which it has been
preserved, it was noted that the alcohol gave off a fairly strong foul odor as
well. Since the production of a foul odor is common with other members of
this section, it is presumed that the new species does the same.

Etymology.—The new species has been named for Mr. Simon Wako who
collected and submitted the specimen to me—along with many other spect-
mens—for identification. Mr. Wako comes from Crater Mountain area where
he and others from that area are involved with the Wild Life Management
project there.

Notes. —Only spirit material of the new species was available and the il-
lustration of B. wakoi (Fig. 1) was drawn from this material. Based on color
photos provided by Mr. Wako and Crater Mountain Project staff, and by
Dr. Geoff Stocker of a flowering specimen in his private collection, the pet-
als and sepal are glossy olive-green with purple to maroon veins and deep
purple to maroon at their base; externally they are darker, almost glossy
brown in color. The labellum is deep glossy red with the two dorsal ridges
paler, almost pink. The column is dorsally white to pink with some red
lines near its base and ventrally deep red. The anther is red and the pedicel
and ovary are green. The new species differs from B. foetidum, in the color
and color patterns of the sepals, petals, and column. The pedicel and ovary
of the latter species is purple black with white spots whilst that of the B.
wakol is green.

ACKNOWLEDGMENTS

I wish to thank Mr. Simon Wako for providing the specimen of B. wakor.
Dr J. Stocker made valuable comments and suggestions during the prepa-
ration of this paper. To Mr. R.J.E Henderson, Queensland Herbarium, Brisbane,
I owe a special thanks for providing assistance with the Latin text.

Technical support for Mr. Wako and other village parabiologists in the
Training Local Observer Program has been provide by the Research and
Conservation Foundation of Papua New Guinea and the Wildlife Conser-
vation with support from the Biodiversity Support Program (BSP) and their
assistance is gratefully acknowledged here. BSP is a Consortium of the World
Wildlife Fund, The Nature Conservancy and the World Resources Insti-
tute, with funding by the United States Agency for International Develop-
ment. The opinions expressed herein are those of the author and do not necessarily
reflect those of the U.S Agency for International Development.

REFERENCES

SCHLECHTER, R. 1911—1914. Die Orchidacee von mein i Guinea. Rep., Beih.1:748.
SMITH, J.J. 1929. Nova Guinea XIV, 475. Plate 7, No.1

1024 Sipa 18(A)
BOOK REVIEW

WILLIAM F. LAURENCE and RICHARD O. BIERREGAARD, Jr. (Editors). 1997.
Tropical Forest Remnants. Ecology, Management and Conservation
of Fragmented Communities. (ISBN 0-226-46898-4, hbk; ISBN
0-226-46899-2, pbk.). The University of Chicago Press, 11030 South
Langley Avenue, Chicago, IL 60628. $27.50. 525 pp. text, 6 pp. con-
tributors, 55 pp. references, 28 pp. indices.

This book contains a series of summary papers from a symposium held in 1995 during the
annual meeting of the Ecological Society of America. While the information may be some-
what dated, the volume is clearly a well-balanced summary of the most important con-
cepts involved with habitat fragments, their restoration and long-term management of the
biota resident therein. Given that tropical habitats are increasingly fragmented, the basic
principles outlined in this book become a pre-requisite element in the formation of every
new conservation biologist. It is important to note that while the papers presented in the
book were written for 1995, they have all been supplied with updated references, so that
frequent references to material published in 1997 is includec

The book is divided in seven sections, including: 1) The scale and economics of tropical
deforestation, 2) physical processes and edge effects, 3) Opies forest faunas, 4) plants and
plant-animal interactions, 5) restoration and man f fragmented landscz apes, 6) site

selection and design of tropical nature reserves, a 7) senate and new perspectives.

Tropical Forest Remnants would be an ideal book for a graduate seminar series. Given that
our planet is largely comprised of a mosaic of remnant landscapes, varying in size, aspect,
slope, hydrology, climate and proximity to other fragmented landscapes, specific training
in fragment dynamics is increasingly important. To introduce the concept of tropical for-
est fragmentation to student conservation biologists, | heartily recommend that papers from
Section VII, summary and new perspectives, be presented first, especially Chapter 4 2, “Tropical

Forest Fragmentation: Synthesis of a Diverse and Dynamic Discipline.” From sheie, read-
ing of the chapters cited therein provide the most useful way to present the copious mate-
rial of this book, the large reference list, and indeed, supplementary material published
since 1997. The book covers, like no other I have seen, a cogent summary of how land-
scapes have become fragmented, how fragmentation among various elements of the biota
should be measured, and what the long-term effects of fragment size, shape, location, etc.
mean to each of those phyla for which we have data. Whether one is strictly concer ened
with tropical phenomena or not, there are valuable lessons for application in understand-
ing habitat fragmentation for other biomes as well. I highly recommend this book to all
who concern themselves with natural history, conservation biology, natural resource man-
agement, ecology and systematic biology.—/John_J. Pipoly HL.

Sipa 18(4): 1024. 1999

NEW COMBINATIONS IN THE
MELASTOMATACEAE FROM HISPANIOLA

HENRI ALAIN LIOGIER
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.

ABSTRACT
Eighteen new combinations are made in the Hispaniolan Melastomataceae, transferring the
species of Ossaea and two species of Clidemia into the genera Henriettea, Leandra, and Sagraea.

RESUMEN

Ca ae a | bi i las Mel Je la Espafiola, transfiriendo

las especies de One y dos especies de Clidemia a los géneros Henriettea, eae y Sagraea.

Several years ago, Walter S. Judd and J. Dean Skean Judd 1986, 1989;
Judd & Skean 1991) pointed in several publications to the genus Ossaea DC.
as an arbitrary group; the genus was originally described as having narrowly
triangular to ovate petals with more or less acute apices. I had already ob-
served that the species considered as belonging to this genus did not show
consistent characters that would help to keep them in the same genus.

As Judd studied the problem further, he concluded that the species of
Ossaea had to be transferred to several other genera, taking into account the
inflorescence position and structure and other characters of generic impor-
tance. He separated the species into several groups belonging to different
genera. After studying the herbarium specimens available and with the type
specimens at hand, Iam now able to place these species in their proper genera:
Clidemia D. Don, Leandra Raddi, and Sagraea DC.; I have also determined
one species in Henriettea DC. Iam also transferring two species of Clidemia
to Sagraea. My work is based on Judd’s and Skean’s publications and con-
clusions. Several species have already been transferred either by Judd or by
his associate, James D. Skean, Jr.

n this paper, Iam transferring these species originally described in Ossaea
from the island of Hispaniola to their proper taxonomic position, prima-
rily according to the inflorescence position and structure, and to the num-
ber of petals in the flowers, in the preparation for my forthcoming treat-
ment of the family Melastomataceae.

Leandra has terminal inflorescences with relatively long calyx-lobe pro-
jections, and 4—6-merous flowers.

Sipa 18(4); 1025-1029. 1999

1026 Stipa 18(4)

Sagraea has lateral inflorescences which may occur below the leaves on former
nodes, shorter calyx-lobes; the 4-merous flowers are solitary or paniculate.

] “1

Clidemia has terminal inflorescences which may “Pp axillary OF |
or occur below the leaves on former nodes, with 4—9-merous flowers And
rounded or retuse petals.

Henriettea has acute to acuminate petals, with lateral inflorescences, the
flowers solitary or in glomerules, 4—G-merous.

I have not been able to see all of the type specimens; some were obvi-
ously lost in the Berlin Herbarium. In some cases, the isotypes are available
in several herbaria.

Henriettea hotteana (Urb. & Ekm.) Alain, comb. nov. Basionym: Ossaea hotteana
Urb. & Ekm., Ark. Bot. 22A(17):58. 1929. Type: HAITI. Depr. Sup-Est: Morne
Delcour, near Pestel, EAman 9008 (HOLOTYPE: S: IsoTyPE: US),

at

This plant has been collected several times by E.L. Ekman in Massif de
la Hotte and near Jérémie, in Haiti. It fits in Hevriettea by its axillary glom-
erate inflorescences and its acuminate petals; it is 4-merous. Several collec-
tions are from the same area as the type collection. It is endemic to Hispaniola.

Leandra humilis (Urb. as Ekm.) Alain, comb. nov. Basionym: Ossaea —
Ekm., Ark. Bot. 22A(17):66. 1929, Type: HAITI: Derr. Sup-Est: Grand
Gosier, Massif de la Selle, Pos 6780 (HOLOTYPE: S; IsoTypE: US).
Ossaea parvifolia Alain, Brittonia 20:158. 1968.
Leandra humilis is a small shrub with terminal solitary sessile 4-merous
flowers, and acuminate petals. It has been collected several times both in
Haiti and in the Dominican Republic. Endemic to Hispaniola.

Leandra ag hae (Urb.) Alain, comb. nov. Basionym: Ossaea hybophylla Utb.,
Ark. Bot. 21A(5):51. 1927. Type: ee Deépr. Centre: Pérodin, Massif des Cahos,
ips 3440 (HOLOTYPE: S; ISOTYPE: NY).
This is a small tree with a terminal inflorescences and 4 acumi-
nate petals. Known only from the type collection. Endemic to Hispaniola.

—— marigotiana (U aD: & Ekm.) Alain, comb. nov. a Osscaea belive
Irb. & Ekm., Ark. Bot. 22A(17):65. 1929. Type: HAITI: Depr. Sup-Est: Marigo
eee de la Selle, Ekman 10071 (HOLOTYPE: S; ISOTYPES: a NY).
This shrub has terminal paniculate inflorescences with 4 acuminate pet-
als. Known only from the type collection. Endemic to Hispaniola.
Leandra ie ha seas ease & Ekm.) Alain, comb. nov. Basionym: Ossaea polychaeta

>, & Ekm., Ark. Bot. 23A(11):27. 1931. Type: DOMINICAN REPUBLIC. Prov. La
VeGa: Loma La Campana, Cordillera Central, EAman 11522 (HOLOTYPE: S; ISOTYPE: NY).

Ly
=
—

Ossaea urbantana Alain, Brittonia 20:58. 1968.

LioGieR, New combinations in the Melastomataceae 1027

A shrub with terminal head-like inflorescnces, the flowers 5-merous with
lanceolate petals. The species has been collected several times in the same
region. Endemic to Hispaniola.

Note.—Urban described two species in Ossaea with quite similar names:
Ossaea polychaeta, now Leandra polychaeta, and Ossaea polychaete now Sagraea
polychaete. This was the reason for a new name published by me as Ossaea
uvbaniana Alain (1968), now obsolete.

Sagraea abbottii (Urb.) Alain, comb. nov. Bastony: Ossaea abbottii Urb., Ark.
t. 22A(17):57. 1929. Type: DOMINICAN REPUBLIC. Prov. Duarre: Loma Quita
Espuela, San Francisco de Macoris, Abbott 182 (HOLOTYPE: ?).
A small tree with axillary, pedunculate head-like inflorescences, 4-merous
flowers and acuminate petals. Known only from the type specimen. Endemic
to Hispaniola.

Sagraea barahonensis (Urb. & Ekm.) Alain, comb. she BASIONYM: ees eae
Urb. & Ekm., Ark. Bot. 22A(17):61. 1929. Typr: DOMINICAN REPUBLIC. P
BaORUCO: Seca de los Comisarios, Ekman 6778 (HOLOTYPE: S).

A 2 m tall shrub with axillary, racemose or paniculate inflorescences, 4-
merous flowers and petals narrowed at apex, not acuminate. Rare in the area.

Endemic to Hispaniola.

oes cinerea (Cogn.) Alain, comb. nov. BasionyM: Ossaea cinerea Cogn., Urb.,
Symb. Ant. 7:316. 1912. Type: HAITI: Depr. GRAND’ Anse: Morne Vandeveld, pling
des Cayes, PE. Christ 1953 (isotype: NY
A shrub with sessile, glomerate, 4-merous flowers; the petals are acute,
not acuminate. It has been collected several times in southern Haiti. En-
demic to Hispaniola.

Sagraea sunalee (Urb. & Ekm.) Alain, comb. nov. BasionyM: Ossaea curvipila
Urb. & ., Ark. Bot. 22A(17):61. 1929. Type: HAITI. Depr. GRAND’ ANSE: Formon
in Massif de | a Hotte, Ekman 7581 (HOLOTYPE: S; ISOTYPE: US).

A shrub up to 2 m tall, with axillary, subcymose, subsessile inflorescences
1—3-flowered, 4-merous flowers. Known from the type collection and from
Formon (Judd 3469, FLAS). Endemic to Hispaniola.

Sagraea ellipsoidea (Urb. & Ekm.) Alain, comb. nov. ay YM: Ossaea ee
Urb. & Ekm., Ark. Bot. 22A(17):63. 1929. Type: HAITI. Depr. GRAND’ ANSE: near
Les Rose Massibae la Hotte, Ekman 10190 (HOLOTYPE: oe ISOTYPE: NY).

A shrub up to 2 m tall with axillary, subsessile cymose inflorescences, 1—
5-flowered; the flowers are 4-merous, the petals ovate. It has also been col-
lected in the Dominican Republic: Prov. Pedernales, Bahoruco Mountains.
Endemic to Hispaniola.

1028 Sipa 18(4)
— fuertesit (Cogn., Urb. Seen ae Nov. BasionyM: Clidemia fuertesii
Cogn., Urb., Symb. Ant. 7:529. . Type: DOMINICAN REPUBLIC. Prov. La
Vek Secabatoa, Central eae lera, pee 1677 UsoryPE: NY).
A slender shrub with axillary |-flowered capillary peduncles, 4-merous
flowers and triangular petals. It is fairly common both in the Central Mountains
and in the Bahoruco Range. Endemic to Hispaniola.

Sagraea gracilis (Alain) Alain, comb. nov. Basionya: Ossaea gracilis Alain, Phytologia
22:167. 1971. Type: DOMINICAN REPUBLIC. Prov. BARAHONA: Monteada Nueva,
el Bahoruco, Alain H. Liogier 11622 (HoLoryrE: NY)

Sierra c

A small shrub, about 1.5 m tall with axillary solitary pedunculate flowers,

these 4-merous, the petals acute. It is fairly common in the area. Endemic
to Hispaniola.

Sagraea lanceifolia (Urb.) Alain, comb. nov. BasionyM: Ossaea lanceifolia Urb.,
Ark. Bot. 22A(17):62. 1929. Type: HAITI. Depr. GRAND’ ANSE: Dame-Marie, Mas-
sif de la Hocte, Ekman 10318 (HOLOTYPE: S),
Shrubby with axillary, subsessile, cymose, 1—3-flowered inflorescences;
the flowers are 4-merous, the petals acuminate. Known only from the type
collection. Endemic to Hispaniola.

ee sacle auies ae ) Alain, comb. nov. Basiony: Clidemia oligantha Urb.,
A(5):45. 1927. Tyrer: HAITI. Derr. Norn: Morne Brigand, Massif du
a aces 2943 (je pe: NY).
A shrub 2—3 m tall with axillary, paniculate, 3—7-flowered inflorescences,
4-merous flowers and ovate, obtuse petals. It has been collected several times,
both in Haiti and in the Dominican Republic. Endemic to Hispaniola.

_

Sagraea polychaete (Urb. & Ekm.) Alain, comb. nov. Basionym: Ossaea poly-
chaete Urb. & Ekm., Ark. Bot. 22A(17):60. 1929. Type: HAITI. Derr. Sup-Esr: near
Jérémie, Massif de a Hotte, Ekman 10401 (HoLoryrt: S).

Shrubby with flowers in axillary sessile groups, 4-merous, the petals rect-
angular, rounded and minutely acuminate. Known only from the type col-
lection. Endemic to Hispaniola.

Sagraea pusilliflora (Cogn., A. & C.DC.) Alain, _— nov. BasionyM: Clidemia
ie Cogn. in A. & C. DC., Mon. Phan. 7:1010. 1891. : DOMINICAN
PUBLIC. Prov. Purrro Piatra: Rio Mameyes, near nee Plata, ae pe (isoryPe: NY).

Asmall shrub about | m tall, with axillary nearly sessile, subsolitary flowers,
these 4-merous. This plant is fairly common, both in Haiti and in the Do-
minican Republic. Endemic to Hispaniola.

Sagraea pas oe ) Alain, comb. nov. Basionym: Ossaea setulosa Urb., Ark.
Bot. 17 1921. Type: HAITI. Derr. Granp’ Anse: Morne La Hotte, ee 157
eee ?).

Liocier, New combinations in the Melastomataceae 1029

A shrub with axillary paniculate few-flowered inflorescences, the flowers
are 4-merous, the petals acute. Another collection from Torbec; also nu-
merous collections from the Formon region, Haiti, by Judd and Skean. Endemic

to Hispaniola.

Sagraea woodsui (Judd & Skean) Alain, comb. nov. Basionym: oes woodsii

udd & Skean, Bull. Florida State Mus. Biol Sci. 32:141. 1987. Type: HAITI: Depr.

GRAND’ ANSE: Morne Formon, Maciel la Hotte, J.D. Skean 1367 aoe FLAS).

A shrub or small tree to 4 m tall, with axillary fasciculate few-flowered

cymes; the flowers are 4-merous, the petals acute. There are numerous col-
lections from the Formon area at FLAS. Endemic to Hispaniola.

NOTE: GENERIC STATUS IS UNCERTAIN
Ossaea rubrinervis Urb. & Ekm., Ark. Bot. 22A(17):59. 1929. Typr: HAITI
Dept. GRAND’ANSE: Les Roseaux, Massif de la Hotte, EAman 10120 (HOLOTYPE: S).
Known only from the type collection which is a sterile specimen.

ACKNOWLEDGMENTS

Tam grateful to the BRIT facilities, both in the Herbarium and in the
Library; to the New York Botanical Garden, to The Rijksherbarium at Stockholm,
and to the Herbarium of the Botanic Garden in Santo Domingo, for the
loan of specimens. And through the years, for the use of the facilities of the
several herbaria visited to study their collections. Iam also grateful to James
D. Skean, Jr. for his revision of this manuscript and his helpful suggestions.

REFERENCES
Jupp, W. S. 1986. Taxonomic studies in the a (Melastomataceae). I. Variation in
inflorescence position. Brittonia 38:150—16
se es OSs Flonistic sae of Morne : Visite and Pic Macaya National Parks,
Haiti. Bull. Florida State Biol. Sci. 32:1—36.
een aaa ones in the Miconteae (Melastomataceae). III. Cladis-
tic aoa of z eee flowered taxa. Ann. Missous Bot. ee 76: 195 bee
-D. SKEAN, JR. 1991. Ti
IV. Generic ee among terminal-flowered taxa. Bull. Florida State Mus., Biol.
Sci. 36:25-84.

1030 Stipa 18(4
BOOK REVIEW

MicuarL A. PoweLt. 1998. Trees and Shrubs of the Trans-Pecos and

Adjacent Areas. Revised Edition. (ISBN 0-292-76579-7, hbk.; ISBN

0-292-7653-8, pbk.). The University of Texas Press, P. O. Box 7819,

Austin, TX 78713-7819. $34.95, pbk; $75.00, hbk. 464 pp., 35 B&

W photos, 370 line drawings.

~S

The Trans-Pecos region of Texas, with its numerous montane habitats and low, arid basins
is substantially different from other regions of the state. The Trans-Pecos is defined geo-
graphically rather than floristically in this work, such that portions Sof the Edwards Plateau
Vegetational Area, and the Chihuahuan Desert are also included. Of course, the Davis Mountains
are also home to a great number of interesting and endemic species. Five general vegeta-
tion types are recognized.

The present revision of the first edition includes taxonomic or nomenclatural updates in

ert

62 genera treated, and new distributional or taxonomic data for another 60. The nomen-
clature and presentation more closely correspond to hierarchies recognized by John Kartesz
and others, with supplements from monographers and other sources. In some significant
cases, keys were re-written. The book covers 447 species of woody plants in 203 genera
and 70 families, and is divided into 6 functional Parts, including che 1) introduction, 2)
key to the classes and subclasses, 3) descriptive flora, 4) selected glossary, 5) literature cited,
and 6) index

Trees and Siu of the Trans-Pecos and Adjacent Areas is very straightforward, easy to use,
and will surely be useful to all who visit, botanize, study, or otherwise witness the beauty
of the Trans-Pecos area of Texas. If we can overlook a few ss pesetting errors Ju N glandaceae
and not Juglandaceae), the book is perfect for a floristic seminar, any nature study tour, or
jusc to have along to botanize. The glossary is concise “and useful, although it is not illus-
trated, the literature cited is extensive, and the index is very helpful and ene to use. As
one might expect, the revision of a classic work has produced something oe R AND

3ETTER, a 4 field guide and yet another volume to fit in the side pocket of one’s car, so

can be drawn more quickly. I recommend it for everyone in the ee Pipoly HHL

Sipa 18(4): 1030. 1999

PENNISETUM ADVENA SP. NOV.
(POACEAE: PANICEAE): A COMMON
ORNAMENTAL GRASS THROUGHOUT
THE SOUTHERN UNITED STATES

JOSEPH K. WIPFF

Pure Seed Testing, Inc.
P.O. Box 449
Hubbard, OR 97032, U.S.A.
joseph @ turf-seed.com
J.E VELDKAMP
Rijksherbarium (L)/ Hortus Botanicus

Box 9514

NL-2300 RA

Leiden, THE NETHERLANDS
J AL h(a

lyhb. leidenuniy, nl

VOEUAREANEL

ABSTRACT

Pennisetum advena Wipftf & Veldkamp, a common ornamental grass in the southern United
States, is described and illustrated. A key is provided to separate it from similar taxa (P.
macrostachys, P. orientale, and P. setaceum). Pennisetum advena is readily distinguished from P.
setaceum, its closest putative relative, by vegetative, inflorescence, fascicle, and spikelet characters.

RESUMEN

e describe e ilustra Pennisetum advena Wipth & Veldkamp, una graminea ornamental
eas enel sur de los Estados Unidos. Se ofrece una clave para separarla de taxa similares
(P. macrostachys, P. orientale, y P. setaceum). Pennisetum advena se distingue facilmente de P
setacewm, la especie mas préxima, por los caracteres vegetativos, de la inflorescencia, del fasciculo

y de la espiguilla.

Pennisetum Rich., a genus of + 80 species, is found in tropical, subtropi-
cal, and temperate regions (Clayton & Renvoize 1986). It occupies a di-
verse range of habitats, including riparian, savanna, desert, forest, and montane.
Of the + 80 species, 40 are known to occur in the New World. Twenty-six
of the 40 are native to México, Central and/or South America, 14 are intro-
duced into the New World. Seventeen species have been introduced into
the United States. This genus contains species that are important as grain
(cereals) [e.g. P. glaucum (L.) R.Br.}, forage {e.g. P. ci/iare (L.) Link, P. flaccidum
Griseb., P. glaucum, P. orientale Willd. ex Rich., and P. purpureum Schumach.],
soil binding and lawns (e.g. P. clandestinum Hochst. ex Chiov.; and orna-

SipA 18(4): 1031-1036. 1999

1032 Sipa 1L8(4)

mentals [e.g. P. advena Wiptf & Veldkamp, P. a/opecuroides (L.) Spreng., P.
macrostachys (Brongn.) Trin., P. orientale, P. purpurenm, P. setaceum (Forssk.)
Chiovy., and P. vi//osum R.Br. ex Fresen.}.

Aposporous apomixis has been reported in Old World species of Pennisetum
[e.g. P. advena (reported under the name P. setacenm purple-type, by Simpson
& Bashaw 1969), P. ciliare, P. flaccidum, P. orientale, P. setaceum, and P. villosum).
Research is still needed to understand the intricate morphological patterns
and relationships, that are complicated by the presence of aposporous apo-
mixis. The genus is morphologically and nomenclaturally complex and is
in need of revision. In some cases the relationship of Pennisetum to allied
genera (e.g. Cenchrus L.) is unresolved.

Since Pennisetum is not native in the United States, it is not well known
to American agrostologists. This, in part, has lead to the overlooking of P.
advena as a distinct species which has become a common ornamental grass
in the United States. This overlooked species will be referred to as P. setacenm
‘Rubrum,’ its current horticultural name.

The first record of P. setacenm ‘“Rubrum’ in the United States was reported
by Hitchcock (1916) as a form of P. rappel7i Steud. (= P. setaceum). Hitchcock
(1916) wrote, “A half-hardy form with dark purplish foliage and purplish
crimson spikes has recently been intro. under the name P. cupreum. It does
not reproduce reliably from seed.” The name P. ‘capream’ is a horticulture
name and was never validly published {see also the index to Bailey (1917),
where P. ‘cuprenm’ is cited as a horticultural name and a form of P. ruppelii J.
The use of P. cupreum Hitche. (or Hitchc. ex L.H. Bailey) is incorrect, since
Hitchcock and other authors have only accepted it as a horticulture name
fora “form” or “horticultural variety” of P. setacenm and not asa validly published
combination (Greuter et al. 1994: Article 34.1). Since its introduction in
L916, P. setaceum ‘Rubrum’ has since become one of the most popular orna-
mental grasses in the United States (Greenlee 1992). Darke (1994) com-
mented that P. setacenm‘Rubrum’ might not belong to P. setaceum.

Simpson and Bashaw (1969) published cytological and reproductive char-
acteristics of P. setaceuam. The two morphological types of plants studied were
designated as “green” or “purple.” The description of the “purple” type appeared
to refer to P. setaceum ‘Rubrum’. Fortunately, Simpson (Texas Agricultural
Experiment Station, Stephenville, Texas) had maintained a clone of this plant
in a greenhouse. In 1987, Kenneth Hignight and the senior author were
able to examine this plant and confirmed that it was P. setaceum ‘Rubrum’.
So began a 10-year search for a valid scientific name for P. setaceum ‘Rubrum’,

In 1987, a specimen of P. setacenm ‘Rubrum’ sent to the Royal Botanical
Gardens (K) was reported as being “similar” or “with affinities” to P. macrostachys,
a robust species from Malesia which also has purple leaves. This research
into P. setaceum ‘Rubrum’ continued as time permitted. In November 1992

WIPFF AND VELDKAMP, Pennisetum advena in the southern US 1033

while working on the treatment of Pennisetum for the forthcoming Manual
of North American Grasses, photographs of the plate of Gymnotrix macrostachys
Brongn. (= Pennisetum macrostachys) in Duperrey (1829) and of the type specimen
from the Muséum National d’Histoire Naturelle (P), were obtained. Upon
examination, it was obvious that P. setacevm ‘Rubrum’ and P. macrostachys
were not the same taxon. Pennisetum macrostachys has only antrorsely scaberulous
bristles in a fascicle, one spikelet per fascicle, and a puberulous inflorescence
axis. Pennisetum setaceum ‘Rubrum’ has two kinds of bristles in a fascicle; an
inner series of plumose, ciliate bristles and an outer series of antrorsely scaberous
bristles, 1-3 spikelets per fascicle and a papillose pubescent infloresence axis.

Germplasm of P. macrostachys from National Germplasm Center in Georgia
was obtained. This accession (PI 354266), originally collected in New Guinea
(Malesia), is actually a green form of P. setacewm ‘Rubrum’. In 1995, the senior
author sent a specimen of P. setaceum ‘Rubrum’ to the junior author, who is
an authority on the grasses of Malesia. The junior author spent two years
searching for the validly published name for this taxon. Also, in 1995, W/.D.
Clayton (K) was contacted for assistance, but he too was not able to puta
name to this mysterious taxon.

After years of unsuccessful searching for a satisfactory identification and
careful examination of the species of Pennisetum known to science, we be-
lieve that this is an undescribed species.

Pennisetum advena Wipff & Veldkamp, sp. as (Fig. 1). Typus: UNITED
STATES. Texas. Brazos Co.: Cultivated at Texas A&M University, College Station,
Texas, commonly used ornamental ai in the area, 18 Sep 1990, Joseph K. Wipff
1723 (HOLOTYPE: L; IsorypEs: K, MO, UTC).

Pennisetum advena a P. setaceo cognatio sua proxima ut videtur facile distinctum in
folii laminis 6-11 mm latis planis costa non-incrassata, culmo in nodis aeriis plerumque
iterum ramoso, oe eee medio involucris 10-17 per sectionem | cm ramo primario
1-2 mm longo, involucro setarum serie interiore setis 8-16 ciliatis vel plumosis, gluma
primaria 0.5—1 mm longa, flore inferiore staminato.

Plants perennial (annual in temperate climates), cespitose, without rhi-
zomes or stolons. Cu/ms 100-150 cm tall, erect; nodes glabrous, usually with
some secondary branching at aerial nodes. Leaves: (measurements taken from

the 2nd and 3rd uppermost leaves); sheaths glabrous, margins ciliate; /igules

0.5—0.8 mm long, a ciliate membrane; b/ades 33-52 cm long, 6-11 mm

wide, flat, burgundy (rarely green), mid-vein not noticeably thickened, margins

antrorsely scaberulous and ciliate at base. Panicles 23—32 cm long, 30-58

mm wide, flexuous and drooping, burgundy (rarely pale or whicish-green);

central axis terete, pubescent with papillose trichomes. Fascicles (Involu-

cres): 10-17 per 1 cm section (mid-inflorescence), with 1—3 spikelets. Pr7-
mary branch \-2 mm long (the length from base of branch to uppermost
bristle (primary bristle). Fascicle stalk (or stipe) 0.5—1.1 mm long {the length

1034 Sipa 18(4)

. Pennisetum advena (Wipff 1723). A. Habit. B. Fascicle. €
one central axis and fascicle arrangement.

. | cm section of panicle

from base of primary branch to first (or lowermost) bristle}. Primary bristle
21.3—33.6 mm long, noticeably longer than the other bristles, papillose
ciliate. Two types of bristles in fascicle: an outer bristles series of 43—68 bristles
1.2-18.5 mm long, antrorsely scaberulous; an ‘nner bristles series of 4-10
bristles, 1 1.7—25 mm long, papillose ciliate. Spzke/ers subsessile or pedicelled
in fascicle. Central Spikelet 5.3-6.5 mm long; pedice/ 0.1—0.3 mm long.
Glumes unequal; first e/ime 0.5—1 mm long, O-veined; second glume 1.9—3.6
mm long, 0—1-veined, about 1/2 as long as spikelet; Lower Floret stami-

nate. First lemma 4.7—-6.1 mm long, 5(—6)-veined. Palea 4.5—5.0 mm long.

WHIPFE AND VELDKAMP, Pennisetum advena in the southern US 1035

Anthers 3, 2-2.5 mm long. Upper Floret: second lemma 5.2-6.1 mm long,
5-veined. Palea 4.7-5.6 mm long, 2-veined. Anthers 3, 2.5—2.7 mm long.
Caryopses infrequently produced in United States plants.

ay, r }
Chr OMLGSIUTTLE FLEEHELUCE

2n = 54 {reported under the name P. setaceum purple-
type (Simpson & Bashaw 1969)}.

Method of reproduction.—aposporous apomixis [reported under the name
P. setaceum purple-type (Simpson & Bashaw 1969)}

Phenology.—May until first freeze.

Distribution. —Cultivated throughout the United States. Will not per-
sist in areas where winter temperatures fall below freezing for any length of
time. In areas with below freezing temperatures, it is used as an annual and
replanted every year or moved into a greenhouse. In southern Florida, Texas
and California, and Hawaii it will persist, but rarely escapes

Etymology.—The specific epithet, advena, is Latin for “visitor” or “stranger”;
referring to its mistaken identity and unknown origin. It is believed to have
originated from the Old World.

KEY TO SIMILAR TAXA OF PENNISETUM

1. Fascicle with two types of bristles: an inner series (bristles closest to spike
let) with bristles long-ciliate, and an outer series of antrorsely scaberulous
bristles. Primary bristle noticeable longer than other bristles in fascicle. Fascicles
with 1-10 spikelets per fascicle. Inflorescence axis pubescen

|. Fascicle with all bristles antrorsely scaberulous. Fascicles with only 1 spike-
let per fascicle. Primary bristle not noticeably longer than other bristles in
en Inflorescence axis puberulous
2. Mid-culm leaf blades (3-93.51

vein not noticeably thickened

P. macrostachys
| mm wide, flat, green or burgundy; mid-

2. Mid-culm leaf blades 2—3.5 mm wide, convolute or folded, green; mid-
vein noticeably thickened
3. Culm nodes pubescent. Plan

P. setaceum
s with rhizomes. Ligule 1-2 mm long.
Fascicles white. nae bristle Sk fascicle with 0—24 terete, scaberulous
bristles. Leaf blades green. Inflorescence erect or arching .............0.. P. orientale
Culm es glabrous. Plants without rhizomes. Ligule less than 1 mm
iene. Fasccles canes (rarely pale green). Outer bristle series of fas-
cicle with 43-58, te ;

jos

scaberulous bristles. Leaf blades burgundy (rarely
green). ee ee and drooping P. advena

Pennisetum advena is readily distinguished from P. setacem, its closest putative
relative, by the following characters. Pennisetum advena: |) leaf blades 6-11
mm wide; flat, mid-vein not thickened; 2) usually with secondary branch-
ing at aerial culm nodes; 3) 10-17 fascicles per 1 cm section (mid- cence);
4) primary branch of fascicles (mid-inflorescence) 1-2 mm ae 5) j inner
bristle series of fascicle with 4—10 ciliate or plumose bristles; 6) first glume
0.5—l mm long; and 7) lower floret staminate. Pennisetum setaceum: 1) leaf
blades 2—3.5 mm wide, convolute, mid-vein conspicuously thickened; 2)

1036 Sipa 18(4)

no secondary branching at aerial culm nodes; 3) 8-10 fascicles per 1 cm
section (mid-inflorescence); 4) primary branch of fascicles (mid-inflorescence)
2.34.5 mm long; 5) inner bristle series of fascicle with 8-16 ciliate or plumose
bristles; 6) first glume absent (rarely present, up to 0.3 mm long); and 7)
lower floret neuter (rarely staminate).
ACKNOWLEDGMENTS

We thank the curators at P for assistance with type specimens (photo-
graphs), the research librarians at BRIT and NY for their support with re-
quests for literature, and W.D. Clayton (K) for his time and assistance. We
thank Stanley D. Jones (BRCH) and Gretchen D. Jones (USDA, AWPMRU)
for their review and suggestions on the manuscripts. The illustration was
prepared by Eddy Dawson.

REFERENCES
BaiLey, L.H. 1917. The standard cyclopedia of horticulture, Vol VI-Z (Vol. 6). The MacMillan
Company, New York
Crayton, W.D. and $.A. RENvoize. 1986. Genera graminum: Grasses of the World. Kew
Bull., Addit. Ser. XIU. Her as Stationery Office; London.
Darke, R. 1994. Manual of the grasses. In: M. Griffiths, ser. ed. The new royal horticul-
tural society dictionary of ae series. Timber ae Sesion ,OR

Duprrrey, LI. 1829. Voyage autour du monde. Phanerogamie. Acdiun enema Paris.

GREENLEE, J. 1992. The encyclopedia of ornamental grasses. Rodale Press, Emmaus, PA.

Greuter, W., ev AL. 1994. Incernational code of botanical momenclature (Tokyo ¢ C ode).
Koeltz Scientific Books, Kénigstein, Germany.

HitcHcock, A.S. 1916. Penvitetum, In: L.H. Bailey. The standard cyclopedia of horticul-
ture. Vol. V-PR (Vol. 5). The MacMillan Company, NY.

Simpson, C.E. and E.C. BasHaw. 1969. Cytology and re ae tive characteristics in Pennisetum
setacenm. Amer. J. Bot. 56:31—36.

NEW COMBINATIONS IN SETARIA
(POACEAE: PANICEAE)

WILLIAM E. FOX, III and STEPHAN L. HATCH

S.M. Tracy Herbarium (TAES)
Department of Rangeland Ecology & Management
Texas AGM University
College Station, TX 77843-2126, U.S.A.

eae! ba) ae
S-hatth@ tami. eau

ABSTRACT

Setaria subgenus Reverchoniae is proposed. New combinations in subgenus Reverchoniae,
Setaria reverchonit (Vasey) Pilg. subsp. ramiseta (Scribn.) W.E. Fox, S. reverchonii (Vasey
subsp. firmula (Hitchc. & Chase) W.E. Fox are proposed. New combinations in cae
Paurochaetinm, S. utowanaea (Scribn.) Pilg. var. ophiticola (Hitchc. & Ekman) W.E. Fox, §
utowanaea (Scribn.) Pilg. var. swbtransiens (Hitche. & Ekman) W.E. Fox
I

é x are also proposed.
K evs Sendar: iting the subge nera. su ds pecies of S

| reverchonit, species of subgenus Pawrochaetium,

el varieties op S. wtowanaea are included.

RESUMEN

Se propone Setaria subgénero Reverchoniae. Son propuestas combinaciones nuevas en el
subgénero Reverchoniae, S. reverchonti (Vasey) Pilg. subsp. ramiseta (Scribn.) W.E. Fox y S.
reverchonit (Wasey) Pilg. su are: (Hitchc. & Chase) W.E. Fox,. Son también propuestas
combinaciones nuevas en el subgénero Pawrochaetium, S. utowanaea (Scribn.) Pilg. var. age ‘ola
(Hitche. & Ekman) W.E. Fox y S. wtowanaea (Scribn.) Pilg. var. swbtransiens (Hitche cman)
W.E. Fox. Se incluyen claves para separar los subgéneros, subespecies de S. relat y las
variedades de S$. wlowanaea.

INTRODUCTION

Setaria P. Beauv. is a cosmopolitan genus important in cultivated crops
{S. italica (L.) P. Beauv.}, perennial forage grasses [S. macrostachya H.B.K.}
and noxious weeds [S. sn (L.) PB. Beauv. }.

Setaria (Poaceae:Paniceae) is one of several genera closely related to Pani-
cum. Tribal and generic arrangement of the species of the Poaceae as classified
by Hiickel (1887) has been revised by Prat (1936), Pilger (1954), Stebbins
(1956), Clayton and Renvoize (1986) and Soderstrom (1986). However, the
circumscription of the Paniceae has remained rather stable. For additional
discussion of previous work see Fox (1999). Most taxa in Sefaria can be eas-
ily differentiated from its closest relatives Panicum and Paspalum by the presence
of bristles subtending the spikelets, these representing modified inflorescence
branches.

Sipa 18(4): 1037-1047. 1999

)38 Stpa 18(4)

Rominger (1962) monographed the species of Setaria from North America
and classified 43 species occurring in three subgenera: Prychophyllum (6 species),
Setaria (27 species) and Paurochaetium (10 species). Twenty-five are native
to North America, ten originate from South America and eight are from
the Old World (Rominger 1962).

The subgenus Pawrochaetium forms an “artificial group” (Rominger 1962)
distinguished from the remainder of the genus by the occurrence of only
one bristle usually below the terminal spikelet of the primary branches. The
subgenus occurs as two separate complexes. The first complex ranges from
southern Florida through the West Indies and into the Yucatan region of
Mexico and Belize. The complex includes Setaria distantiflora (A. Richard)
Pilg., 8S. pradana (Léon) Léon, S. leonis (Ekman) Léon, S. ophiticola (Hitche.
& Ekman) Léon, S. swbtransiens Hitche. & Ekman, S. “towanaea (Scribn.) Pilg.,
and S$. chapmanii (Vasey) Pilg.. The second complex occurs from northeast-
ern Mexico through the western two-thirds of Texas, into southern Okla-
homa and west into New Mexico. It extends from the Balcones Escarpment
of Texas to Chaves County, New Mexico. The complex includes S. firmula
(Hitche. & Chase) Pilg., 8. ramiseta (Scribn.) Pilg. and S. reverchonii (Vasey)
Pilg.. Not included in Rominger (1962) is the species 8. variifolia (Swallen)
G. Davidse found in the Yucatan Peninsula of Mexico and south

Rominger (1962) separated the subgenus from the remainder of Setaria
by the presence of a single bristle “usually” below the cerminal spikelet of
each primary branch. Gould (1975) used the same character to separate the
species that occur in Texas; however, he reported a problem with the identification
of specimens based on this character. These problems led to the current research
in the subgenus.

The objectives of the research were to 1) determine the relationships of
the taxa in the Texas/Mexico/Oklahoma/New Mexico complex and 2) pro-
vide a taxonomic treatment of all taxa recognized by Rominger (1962) in
the subgenus Pawrochaetium and S. variifolia.

MATERIALS AND METHODS

Field collections were made during the flowering periods of the species
throughout Texas, Florida and Belize. To insure isolation between the populations,
a minimum distance of five miles was traveled between successive collec-
tion sites. Ten independent specimens from each site were semi-randomly
collected and pressed; selection was biased in favor of mature plants that
did not show any signs of damage from insects, herbivores, trampling, etc.
The West Indies species were studied from herbaria specimens. In some cases
sufficient samples were available of a single collection to serve as a “real”
population. In others, “artificial” populations were created based upon similar
geographic location. A total of 78 populations were studied for the subgenus.

Fox AND HatrcH, New combinations in Setaria 1039

Table 1 lists the 52 characters measured from representative populations
to determine the relationship between the 11 species. All characters of se-
lected populations were measured and statistically analyzed to determine
significant characters for the detailed study of the subgeneric relationships.
Nineteen characters were determined to be significant and included in that
analysis. The following hypothesis was tested: taxa of subgenus Pawrochaetinm
should be classified in one subgenus versus the need to describe a new sub-
genus. Two hundred and ninety two operational taxonomic units (OTU’s),
representing all species included in the study, were measured.

nivariate statistics (mean, standard deviation and range) were obtained
using the PSI-Plot software package (Poly Software International 1996).
Multivariate statistics of principal component analysis (PCA) was obtained
using the NT-SYS software package (Rohlf 1990). Principal components
were derived using correlation matrices.

RESULTS AND DISCUSSION
SUBGENERIC STUDY

Based upon the analysis of the 11 species classified in Searia subgenus
Paurochaetinm, two distinctly separate entities were discovered. These dif-
erences matched the two complexes within subgenus Paurochaetium men-
tioned earlier. Figure | illustrates the separation between the two complexes.
Based upon the relationships revealed through the PCA, geographical
separation, consistent morphological differences and similar habitat requirements

of the complexes, the subgenus Reverchoniae is proposed
Setaria subgenus Reverchoniae W.E. Fox, subgenus nov. BasionyM: Panicum
ee Vasey, Bull. U.S. Dept. Agric. Div. Bot. 8:25. 1889. Type: Reverchon 5.n.

yeus: Setaria reverchonii (Vasey) Pilg.

Setavia oo Ret oo ab subgeneri Pawrochaetio ia a paleae flosculi
inferior, paniculam erectam, spiculam grandiorem (2.1—4.5 mm logan, 1.2—2.6 mm latam)
fortuito dispositam ee ose: atque axe a. scabro on

Setaria subgenus Reverchoniae differs from subgenus Pawrochaetium in the
absence of a palea of the lower floret, erect panicle, larger spikelets (2.1—4.5
mm long, 1.2—2.6 mm wide) that are randomly disposed (not distichous),
and the scabrous axis of the inflorescence.

Setavia sabgenus Reverchoniae contains three taxa previously classified in
the subgenus Paurochaetium (Rominger 1962) including the proposed sub-
species: Setaria reverchonii (Vasey) Pilg. subsp. reverchoni, S. reverchonii (Vasey)
Pilg. subsp. ramiseta (Scribn.) WE. Fox and S, reverchonii (Vasey) Pilg. subsp.
firmula (Hitchc. & Chase) W.E. Fox. Setaria variifolia was included in the
subgenus, but was not treated originally by Rominger (1962). The lack of
a palea in the lower floret best circumscribes the subgenus Reverchoniae

1040 SIpA 18(4)

TABLE |. Fifty-ewo characters used to assess morphological and reproductive variation in Setaria sub-

genus Panrochaetium. All characters were measured for quantitative variation except those labeled as
TS {ewo-state (binary)] and MS [multi-stace qualitative]. Italicized characters were found as significant

and eds in oie final analysis.

Vegetative: growth habit (MS), calm height, culms branched/unbranched (BS), internodes hollow/solid
(BS), node pubescence present/absent , leaves basal/throughout (BS), leaf sheath pubescence present/
absent (BS), leaf sheath pubescence bank (if pre sent), leaf length, leaf width (widest point), leaf base
width, leaf involled/not tnrolled (BS), leaf pube SCENCE ne absent (BS), leaf pubescence adaxial

present/absent (BS), leaf pubescence margin p nant ime (BS), ‘eallae pubescence present/absent (BS),
collar pubescence length (if present), auriculate trichomes present/absent (BS), avriculate pubescence
length Gf presence), ligule type (MS), /eale length

js)

Inf
lower primary branches, distance between upper primary branches, terminal bristle length lowest

orescence: inflorescence length, inflorescence axis pubescence present/absent (BS), distance between
branch, terminal bristle length upper primary branches, terminal bristle length terminal primary
branch, bristle exceeding spikelet lowest branch (BS), bristle exceeding spikelet upper branch (BS),
bristle exceeding spikelet terminal branch (BS)

Spikelet: spikelet length, sprkelet width, lower aie clasping | (BS), lower glume apex shape, lower glume # of

veins (MS), dower glume length, upper glume equal/not equal fertile lemma, apper glame length, lower glume
# veins (MS), sterile lemma equal/noct equal fertile lemma ae werile lenma length, scerile lemma # veins
(MS), sterile ere present/absent (BS), ster//e Le length, tertile lemma rugose (BS), fertile lemma length,
fertile palea length, caryopsis length, ee width, filament le sie anther length, anther width

(exception S. variifolia that shares other characters aligning it with the sub-
genus). None of the members of subgenus Reverchoniae are sympatric with
subgenus Pawrochaetium. For further detail of the relationships of the two

subgenera refer to Fox (1999).

KEY TO SUBGENERA OF SETARIA

1. Bristles one to many below each spikelet (some without bristles); leaf blades
plicate 2

2. Leaf blades plicate; bristles present below only some of the spikelets
Ptyc hophy Tlum

2. eet blades not piicate, bristles below all spikelets (rarely missing) Setaria
LB only below the terminal spikelet of branch as an extension
of iseaniels leat bl ve not plicate 5

3. Panicles nodding (except S. pradana), bearing remote, appressed, mostly

racemose branches with spikelets two ranked on an undulating axis; cen-

tral inflorescence axis glabrous; palea of lower florets present and conspicuous
Paurochaetium
anicles erect; spikelets arranged randomly on branch; central inflorescence

_P

axis scabrous; palea of lower florets absent (except S. variifolia) ......... Reverchoniae

SUBGENUS REVERCHONIAE

Four taxa are classified in the subgenus Reverchoniae: Setaria reverchonii subsp
reverchonit, S. reverchonit subsp. ramiseta, S. reverchonii subsp. firmula and S.
variifolia. Figure 2 illustrates the relationship of the taxa in Reverchoniae.

Fox AND Harcu, New combinations in Setaria

PC 2
a Florida/West Indian Complex Mil
ioe American Mainland Complex @ ¢€
Fic. 1. Two-dimensional representation of all populations measured for the genus Setaria

subgenus Paurochaetinm (Rominger 1962)

Rominger (1962) classified S. reverchoniz, S. ramiseta, and S. firmila as separate
species following the combinations proposed by Pilger (1940). Other au-
thors who have classified these taxa as species include Hitchcock (1935),
Silveus (1942), Hitchcock (1951) and Gould (1975). When using the keys
prepared by these authors, it becomes evident that the characters used do
not result in consistent identifications. This problem resulted in the study
of this complex and the following combinations.
ae oe g. subsp. ramiseta (Scribn, ) W.E. Fox, comb.

BASIONY : Panicum 3 bacae Scribn., Circ. U.S. Dept. Agric. Agrost. 27:9.
ie Setaria ramisela (Scribn.) Pilg., Engler & Prantl, Die Nat. Pflanzenf. 14e:72.
1940. Type: G. Nealley s.n. oe eee US!, by Rominger 1962).

Panicum gna eyes vee), Bull. U.S. Dept. Bot. 8:25. 1889, non Panicum subspicatum

. Sci. Phys. Nat. 84. 1831.

ee references and illustrations. —Hitchock & Chase, Contr. U.S. Natl.

Herb. 15:24, fig. 5. 1910; Hitchcock, Man. Grasses U.S. 611, fig. 1271. 1935.

Setaria reverchonii (Vasey) Pilg. subsp. firmula (Hitchc. & Chase) W.E.
ox, comb. nov. BasionyM: Panicum firmulum Hitchc. & Chase, Contr. U.S. Natl.

Heck, 15:27, fig. 9. 19LO. Sa (Hitche. & Chase) Pilg. in Engler & Prancl,

Die Nat. Pilauzent 14e:72. 1940. Type: D. Griffiths 6446 (HOLOTYPE:

oi
2

Additional references and ‘ieee ee es ocal reference and illustra-
tion: Hitchcock, Man. Grasses U.S. 612, fig. 1273. 1935.

1042 SipA 18(4)
awa

A

- Aad

A tes, . rf.

a ‘Dpalg Be

e oe

PC 2 bg
a
* e a
r *
*
*
s I bsp. reverchonia fi
S } t ramiseta
f
Setaria reverchonia subsp, firmula e
Ld
Setaria variifolia *

Fic. 2. Two-dimensional representation 1
in Setaria subgenus Reverchoniae.

lustrating the relationships of the taxa described

Setaria reverchonii subsp. ramiseta can be separated from the other taxa of
the subgenus Reverchoniae with a shorter culm length, shorter penultimate
leaf length, narrower leaf width, shorter second glume, shorter upper and
lower lemma and shorter palea of the upper floret. Setaria reverchonii subsp.
reverchonii is separated from subsp. firmu/a by a longer, narrower and usu-
ally involute leaf blade, a narrower leaf blade base above the collar and a
longer lemma of the lower floret. Further references to the subspecies can

be found in Fox (1999),

KEY TO SUBGENUS REVERCHONIAE

1. Palea of lower florets present and well developed; palea of u per (fertile) florets

3.0—3.1 mm long; distribution Yucatan peninsula of ee and Central

America S. variifolia
l

. Palea of lower florets absent or rudimentary; palea of upper florets (1.0—)1.8—
>) mm long; distribution Texas, New Mexico, Oklahoma and north-
ern Mexico
2. Spikelets (2.4—)2.7—2.8(-3. =e mm long, (1.1—)1.5—1.6(—-1.9) mm wide;
second glumes (1.9—)2.4—2.5(—
>

.1) mm long; lemma of lower florets (1.0)

2.3-2.5(—3.2) mm long; on of upper florets (1 .0-)2.2-2.3(-2.9) mm
long; penultimate leaf blades (3.0-)6.8—7 .6(-1 2.1) cm long; panicles (3.7
17.7—20.7(—38.1) cm long

S. reverchonii subsp. ramiseta
. Spikelets (2.9- )3, 2—3.4(—4.0) mm long, (1.2-)1.8—2.0(—3.6) mm wide;
second glumes (2.4—)2.8—3.0(—3.6) mm long; lemma of hls florets (1.4)

i)

2.8—3.0(—3.7) mm long; lemma of upper florets (2.2—)2.7—2.9(—3.4) mm
ees a leaf blades (3.6-)7.9-13.3(— oe. 6) cm ce panicles (9.5)
25.3-32.6(—64.8) cm long 3

Fox AND Hatcu, New combinations in Setaria 1043

3, ponultnat leaf blades (3.6-)11.7—13.3(—28.6) cm long, (1.4—)2.1—2.3(—
wide, involute; leaf blade oe above collar (0.7—)1.3—1.5(-2

mm wide; lemma of lower florets (1.4—)2.9—3.0(—3.7) mm long ...... S. reverchonii
subsp. reverchonii

3. Penultimate leaf blades (5.1—)7.9-8.8(-15.4) cm long, (2.7—)4.4-4.7(-—

9.2) mm wide, flattened; leaf bases above collar (1.4—)3.0—3.2(-5.3) mm
wide; lemma of lower florets (1.9—)2.8—2.9(—3.2) mm long ............. S. reverchonii
subsp. firmula

SUBGENUS PAUROCHAETIUM

The remaining seven taxa are retained the subgenus Pawrochaetium as described
by Rominger (1962). These include the original species of Setaria distantiflora,
S. leonis, S. ophiticola, S. pradana, S. subtransiens, 8. utowanaea and S. chapmanit.
Taxonomic interpretation was difficult in the subgenus Pawrochaetium due
to a small sample size. Unfortunately, few specimens have been collected
over time and available for the study. Hitchcock (1936) and Rominger (1962)
classified S, wtowanaea, S. ophiticola and S. subtransiens as distinct species. However,
based upon the specimens examined in this study the following combina-
tions are proposed.

Setaria utowanaea (Scribn.) Pilg. var. gees (Léon) W.E. Fox, comb.
nov. BASIONYM: Panicum opbiticola Hitchc. & Ekman ex Hitchcock, U.S. Dept. Agric.
ee Publ. 243:293, fig. 282. 1936. oe ophiticola (Hitchc. & Ekman) Léon in

|. Cuba 163. 1946. ae E.L. Ekman 12712 (Ho.oryee: US!).

Setaria utowanaea (Scribn.) Pilg. var. subtransiens (Hitchc. & a!

ae Fox, comb. nov. Basionym: Panicum subtransiens Hitche. & Ekman ex Hitchcock,

. Dept. Agric. Misc. Publ. 243:351, fig. 323. 1936. Type: E.L. Ekman 168 28
ee US!).

Based upon the mid-leaf width, leaf base width, spikelet length, lower
lemma length and palea length of the upper floret, the varieties of Setaria
ntowanaea are classified separately from the remainder of subgenus Pasrochaetinm.
Figure 3 illustrates the relationship of the three taxa in the Sefaria utowanaed
complex. Setaria utowanaea var. subtransiens differs from the other members
of the species witha single bristle present below most spikelets on the branch,
bristle length exceeding the spikelet, and spikelet width. Setaria utowanaea
vat. utowanaea is separated from S. wtowanaea var. ophiticola by blade length,
spikelet width, lack of tufted basal leaves and the presence of involute leaves
on the upper culm. Setaria utowanaea var. ophiticola has shorter leaf blades,
narrower spikelets, a tufted lower leaf arrangement and upper leaves that
are slightly folded or flat. The following key separates the species of subge-
nus Paurochaetinum. A key to separate the varieties of Setaria utowanaea will
follow. Further reference to the taxa of subgenus Pawrochaetium can be found

in Fox (1999).

1044 Stipa 18(4)

A
A
A A
- ®
A i { ] a
A
A A
PC 2 : . .
A @ a ia Ba
Ma a A A
A
Selaria ufowanaea Var. ulowanaca x
A
A Setaria utowanaea Vat. ophiticola o
Setaria utowanaed var. subtransiens Wh
Fic. 3. Two-dimensional representation of the relationship between Setaria utowanaea var.

utowanaed, S, utowanaed vat. ophiticola and S. utowanaea var. subtransiens.

KEY TO SUBGENUS PAUROCHAETIUM

1. Spikelets (1.8—)2.0—2.2(—2.4) mm long, (1.0—)1.1—1.2(-1. Ce wide, ovate,
lacking palea of ae Horets; penultimate - blades (9.3-)15.1-19.1(—34.4)
cm ee (2.3—)3.2—3.9(-7.7) mm wide; leaf blade bases | collar ( -
1.2—2.7(-6.0) mm wide: second glumes subequal to equal fertile lemmas

. chapmanii

Spikelets (1.3—)1.4—2.4(-2.5) mm long, paleas of lower floret conspicuously

present; leaf blade bases above collar (0.3—)0.4—1.4(—1.8) mm wide; second

glumes conspicuously shorter than f te lemmas 2
2. Panicles erect; spikelets (1.7—)1.8—2.0(-2.5) mm long eee se
lemmas (1.4—)1.6—1.8(—2 1 mm longs pie paleas (0.4—)0.8—1.1(—1.4)

mm long; upper paleas (1.2—)1.3—1.5( A mm long S. pradana
. Panicles seen sikees (1.8—-)2.2—2.4(—2.5) mm long [except S.
pe eee —)1.4 —1.7)}, not eae lower lemmas (1.6—)1.9—
2.2(—2.4) mm ong on paleas (0.8—)1.2—1.8(—1.9) mm long; upper paleas

(1.4-)1.6-1. ee 1) mm He 3
3. Spikelets 3-)I 4-1,5(-1.7) mm ee o glumes (0.4—)0.6—0.7(—
0.9) mm lin second ee ).6-)0.9-1.1(-1.3) mm long; cena
(1.1—-)1.3—1.4(-1.5) mm long; len of lower florets (0.3—)0.9-1.1(—
1.3) mm long; palea of upper florets (0.9-)1.0-1.1(-1.3) mm long

S. distantiflora

=

anor

Na)

3. sy aan (1.6—)1.8—2.4(—2.5) mm long; first glumes (0.6—-)0.7—1.3
nm long; cecund glumes (0.7—)1.2—1.7(—2.1) mm long; palea of
lower florets (1.2-)1.6—2.2(—2.4) mm long; palea of upper florets (1.2—)
1.4-1.9 (-2.1) mm long 4
4. Penultimate leaf blades (3.6—)4.7—5.9(—7.4) mm wide: leaf blade base
above collar (1.1—)1.3-1.5(-1.8) mm wide; spikelet (1.6—)1.8—2.0(—

—

Fox AND Hatcu, New combinations in Setaria L045

2.3)mm long; lemma of ae ae —)1.6-1.8(—2.2) mm long;
hale of upper florets (1.2 ae a 7)mm ihe S. leonis
4, Penultimate ae blades (0.9—)1.1—1.3(4. ‘aa wide; leat blade above
collar 0.4-1.3 4) mm wide; ae ets (1.8—)2.2—2.4(—2.5) mm long;
lemma of ie cae (1.6-)1.9-2.2(—2.4) mm long; } ale of upper
florets (1.4—-)1.6-1.9(-2.1) mm long S. utowanaea

KEY TO VARIETIES OF SETARIA UTOWANAEA

—_

. Bristles present below most spikelets on branch, (2.5—)4.0—5.1(—6.4) mm

long, exceeding spikelet S. utowanaea var. subtransiens
1. Bristles present below only terminal spikelet of branch, (0.4—)0.5—3.3(—3.9)
mm long, not exceeding _ : Z
2. Leaf blades (7.8—)13.1-16.0(—-21.9) mm long; — 0.7—0.8(-0.9) mm
wide; basal leaves not ae. upper leaves inve . utowanaea
var. utowanaea
2. Leaf blades 3.3-5.4(-—6.2) mm long; spikelets 0.5—0.7 mm wide; basal
leaves tufted, upper leaves flat or slightly ane toward apex ........ S. utowanaea

var. ophiticola

CONCLUSIONS

Rominger (1962) stated that the separate classification of Setaria subge-
nus Paurochaetium is “artificial.” However, based upon this study and input
from various others, it is our opinion that, although “artificial” by definition,
the eleven taxa form a distinct group distinguishable from the remainder
of Setaria. Based upon this analysis, keys have been developed that consis-
tently separate the taxa from the subgenera Setaria and Ptychophyllum.

However, our understanding of the relationships within what was origi-
nally classified as Setaria subgenus Pawrochaetinm have changed based upon
these analyses. The proposal of the new ibmen: us, S etaria subgenus Reverchoniae,
provides a natural separation of the original taxa recognized by Rominger
(1962). With the proposal of the new subgenus, Setaria now has four recog-
nized subgenera.

The inclusion of Setaria variifolia with the remainder of Setaria subgenus
Reverchoniae added a little known taxon to the subgenera of Setaria that has
a single bristle usually only below the terminal spikelet of the branch. Davidse
(1981) suggested that S. variifolia was closely related to taxa of subgenus
Paurochaetium as defined by Rominger (1962). This taxon provides a chal-
lenge to its classification due to the overlapping characteristics with sev-
eral of the subgenera. However, based upon these analyses, the taxon is best
classified with the subgenus Reverchoniae. Setaria variifolia warrants further
study to determine if this classification is truly defined. A comparative analysis
of all of the subgenera would determine if the classification is correct.

Within Setaria subgenus Reverchoniae two name combinations have been
proposed. Based upon numerical study of morphology, leaf anatomy and

1046 Sipa 18(A4)

DNA content analyses S. reverchonii eas firmula and S. reverchonii subsp
ramiseta have been proposed (Fox 1999). The taxa of the subgenus (exclud-
ing S. varitfolia) show a substantial amount of intergradation,

The remainder of the species originally classified by Rominger (1962)
are retained in the subgenus Pawrochaetinm. However, name combinations
have been proposed for two of the taxa, S. wtowanaea var. ophiticola and S.
utowanaea var. subtransiens.

There is a need for the study of the reproductive behavior of the three
subspecies of 8. reverchonii. The results of such research would help better
understand the relationship of the taxa. If these taxa do not hybridize, it
may indicate a trend towards speciation.

ACKNOWLEDGMENTS

We are grateful to Stanley D. Jones (BRCH) and Gretchen D. Jones (USDA,
AWPMRU) for reviewing the manuscript. We would also like to thank Paul
Fryxell for the Latin translation of subgenus Reverchoniae and Jesus Valdes
Renya for the Spanish translation of the abstract. Thanks is also given to
the following herbaria for loans of specimens used in the study: ARIZ, BRIT,
ENCB, F, FLAS, FTG, JBSD, LL, MO, MICH, NMC, NY, OKL, OKLA,
PAUH, TAES, TEX, UPR, UPRRP, US and USF.

REFERENCES
Crayton, W.D. and S.A. Renvoize. 1986. Genera graminum: Grasses of the world. Her
Majesty's ice Office. London.
Davipsr, G. 1981. Setarta variifolia (Swalle ae s ee a new combination (Gramineae:
Panicoidese) ree Missouri Bot. Gard. 68
Fox, W.E. 1999. A biosystematic study of hey genus Setaria subgenus Paurochaetium and
y ee subgenus Reverchonia. Ph.D. Dissertation. Texas A&M University,

—

the new
College Sta
Gou Lb, FW. i O75, The grasses of Texas. Texas A&M University Press, College Station,
HAckei, E. 1887. Echte Graser. In: Engler & Prantl, Die Natiirlichen Pflanzenfamilien.
e404 pa ee
Hircncock, A.S. 1935. Manual of the grasses of the United States. U.S. Dept. Agric. Misc.
Publ. 200. U.S. Government Printing Office, Washington, D.C.
Hircncock, A.S. 1936. Manual of the grasses of the West Indies. U. S. Dept. Agric. Misc.
Publ. 243. ae Mee sae Printing Office, Washington, D.C
Hircncock, ee 51. Manual of the grasses of the United States. pu Ed. U.S. Dept. Agric.
Publ. ).U. . ee Office, Washington, D.C.
PILGER, . i 40. Granimeae III: Unterfamilie Pancoideae. In: Engler & Prantl Die. Nat.
Pflanzen. 143:70—73
PirGer, R. 1954. Das system der Gramineae. Bot. Jahresb. 76:281—384
Prat, H. 1936. La systématiques des Graminées. Ann. Sci. Nat. Bot., See 10, 18:165—

258.
Poy Sorrware INTERNATIONAL. 1996. Psi—Plot Ver. 5.02, Salt Lake City, Utah.

Fox AND Hatrcu, New combinations in Setaria 1047

ROLE! ay ieee NTSYS-—pe: ees taxonomy and multivariate analysis system, Ver.
1.§ er Software, New York.

re I M. 1962. as of Setaria Sie aca in North America. Illinois Biol.

29. University of Illinois Press, Urb

poe WA. 1942. Grasses: classification and descr siption of See re ia and Pani-
cum in the United States. Published by author, San Antonio, Tex

SODERSTROM, T.R.; K.W. Hitu; C.S. CAMPBELL, and M.E. nya (eds.) 1986. Grass
systematics afd, evolution. Smithsonian Institution Dec Washington, D.C.

EN G.L. 1956. Cytogenetics and evolution of the grass family. Amer. J. Bot. 43:890—

48 Sipa 18(4)
BOOK REVIEW

LAURENCE C, WALKER 1996. Forests: A Naturalist’s Guide to Woodland
Trees. (ISBN 0-292-79112-7, pbk.). The University of Texas Press,
P. O. Box 7819, Austin, TX 78713-7819. $19.95. 302 pp., 68 B &
W photos, 12 figures, 3 tables.

[his book is aimed at the amateur naturalist, to introduce the major groups of timber trees,
their uses, and the forests they grow in. While | a difficulty with terms such as “male
Howers” of Spruce and Fir (microsporangiate strobili), | understand that the author is pur-
posely trying to keep as many of the species descriptions parallel throughout the text, which
he accomplishes. The text is filled with interesting trivia, impor tant general

ical c neepts

important dendrological and morphological features of the trees, many of their uses an id
distribution. It is copiously illustrated and extremely easy to “leaf” through. There are numerous
editorial comments, akin to those sometimes made by senior distinguished lecturers, but
they do not seriously affect the delivery of important information in the text. It is obvious
that Walker is a pragmatic practitioner, and while the theoretical concepts may be slightly
out of date, the general summary information for each group of species is very useful, par-
ticularly to those unfamiliar with which species are commercial, where they grow, and what
is important about

Perhaps because am not a forester, I found such terms as “stink-bomb tree” for Gineko
amusing, as [ dic discussion of its “leaf spigots” and “living scissors” (Chapter 28).
While the ieee may be “folksy” and very informal, it is aptly suited to the intended
audience ee painlessly injects some biological knowledge directly into the reader. The

rojects for the Amateur Naturalist” section at the end of every chapter are cleaver exer-

cises that oe the audience how foresters do their work. While most are very straightfor-
ward, they are all time- and resource-consuming, but again, most things worth doing are.
T ul.

In summary, | recommend this book tor every i kyard naturalist who is curious to know

re reading lists and glossary are extremely us

how the forestry community views the most important kinds of timber trees and their
environments.—John_J. Pipoly LL.

Sipa 18(4): 1048. 1999

THE TAXONOMY OF CYPERUS VIRENS AND
CYPERUS DRUMMONDII (CYPERACEAE) IN THE
SOUTHEASTERN UNITED STATES

RICHARD CARTER and DAVID K. ALEXANDER
Herbarium, Biology Department
Valdosta State University
Valdosta, GA 31698-0015, U.S.A.

CHARLES T. BRYSON

USDA, ARS
Southern Weed Science Research Unit
P.O. Box 350
Stoneville, MS 387706, U.S.A.

ANDREAS LAZARI

Mathematics and Computer Science Department
Valdosta State University
Valdosta, GA 31098, U.S.A.

ABSTRACT

Data for 20 morphological characters taken from recent collections of eG ae virens
group from the southeastern United States were statistically analyzed using }

analysis and discriminant analysis. Based upon results of the analyses one recent ae ob-
servations, it 1s concluded that C. drammondit and C. virens are distinct at the rank of spe-

cies. The taxonomy is revised accordingly. A taxonomic key, technical descriptions, distri-
butional, and ecological data on the two species are presented.

RESUMEN

Los datos de 20 caracteres morfol6gicos de colecciones recientes del grupo de Cyperus

virens del sureste de los Estados Unidos fueron analizados estadfsticamente usando el andlisis

de SOR ROR ON : piney el andlisis discriminante. En base a los resultados de los andalisis
y de

vaciones de campo, se concluye que C. drammondii y C. virens son distintas

en el rango de especie. Se revisan - aeuereo con esto la taxonomia. Se a tear una clave
ela dist ac]

y sob

taxonom ica, desc ripciones las dos espec ies.

INTRODUCTION

Cyperus virens and C. drummondii are closely related and belong to section
Luzuloidei and are characterized by persistent rachillas; bicarinate, decidu-
ous scales; and 1—2 stamens per floret (Kiikenthal 1935-1936; Denton 1978).

Stipa 18(4): 1049-1063. 1999

1050 Stipa 18(A4)

Both taxa have sharply three-angled (triquetrous) culms, which, in combi-
nation with the characteristics listed above, easily separates them from all
other Cyperus species in the southeastern United States. Carter (1990) pro-
vides a key separating C. virens and C. drummondti from other members of

section Luzuloider.

Cyperus virens Michx. has historically presented a taxonomic problem because
of its complex pattern of variation and wide distribution. According to Denton
(1978), C. virens consists of four varieties: C. virens var. virens, C. virens var.
drummondi (Yorr. & Hook.) Kiikenthal, C. virens var. minarum (Boeck.) Denton,
and C. virens var. montanus (Boeck.) Denton. Of these taxa, only C. drummondii
and C. virens var. virens occur in the United States. Cyperus drummondii was
first described in 1836 based upon a specimen collected in Texas by Tho-
mas Drummond (Torrey 1836). Cyperus drummondit has been treated vari-
ously since.

Chapman (1889), like Torrey (1836), recognized Cyperus drummondit.
Subsequently, several floristic manuals include only C. vzrens with no men-
tion of C. drummondii as either a taxon or a synonym (Chapman 1897; Mohr
1901; Small 1933; Correll & Johnston 1970; Correll & Correll 1975; Godfrey
1979: Wunderlin 1982, 1998; Clewell 1985; Thomas & Allen 1993). Hatch
et al. (1990) did not recognize C. drummondii at any rank, while Tucker (1994)
treated it as a synonym of C. virens. Cyperus drummondii was first treated as a
variety of C. virens by Kiikenthal (1935-1936); this view was upheld by
McGivney (1938), Denton (1978), and Adams (1994). Koyama (1970) treated
C. drummondii as a subspecies of C. virens. Other recent authors (Bryson &
Carter 1994; Jones et al. 1997) recognized C. drummondii at the rank of species.

Cyperus virens and C. drummondii are widely distributed in the New World
and are sympatric in the coastal plain of the southeastern United States.
Although C. drummondii has been known from Louisiana and Texas (Torrey
1836; McGivney 1938), it is infrequently collected elsewhere in the south-
eastern United States. Recently, it was reported new from Mississippi (Bryson
& Carter 1994). Kiikenthal (1935—1936) and Denton (1978) documented
the distribution of C. drummondii outside the United States in Nicaragua,
Jamaica, Surinam, Ecuador (Galapagos Islands), and Brazil.

MATERIALS AND METHODS

We examined 57 specimens of C. virens and 44 specimens of C. drummondii.
Specimens were reproductively mature and from the southeastern United
States. Initially, each specimen was identified by a unique number and af-
ter preliminary examination was assigned to C. drummondii or C. virens.

After a survey of the critical literature (Kiikenthal 1935-1936; Denton
1978) anda preliminary study of our collections 20 characters were selected

CARTER ET AL., Taxonomy of Cyperus virens and drummondii 1051

Taste 1. List of characters used in analysis.

PPEDN number of primary peduncles

BRACNL number of primary infloresence bracts exceeding longest ray
CULML culm length (cm

BRACTL maximum primary inflorescence bract length (cm)

RAYL maximum ray length (cm

INF maximum inflorescence width (cm)

SPKW maximum spike width (c

CULMW mid-culm width (mm)

BRACTW maximum mid-bract width (mm)

LEAFW maximum mid-leaf width (mm)

SPKLTW spikelet width (mm)

SCALL scale lengtl )

ACHNL achene length (mm)

ACHNW achene width (mm)

ANTHI anther length (mm)

SCALN scale number per spi t

CIRAT ratio of culm length to inflorescence width

ASRAT ratio of achene length to scale length

ACHLW ratio of achene length to width

BCRAT ratio of longest primary inflorescence bract to culm length

—

for analysis. The characters are listed in Table 1, and all character states are
continuous quantitative. These data were subjected to principal components
analysis (PCA) and discriminant analysis (DA) using Minitab© release 11.21
(Sneath & Sokal 1973; Seber 1984; Anonymous 1996). Data points plotted
using PCA were identified by specimen number and their @ priori species
categories, and eigenvectors generated from a PCA were used to determine
which characters most accounted for the variance observed. All characters
were analyzed using DA to determine order of reliability of characters and
which minimal combination of characters would allow classification of our
specimens into the taxonomic groups defined by PCA

RESULTS

Results of PCA, shown in Table 2 and Figure 1, indicate the specimens form
discrete clusters corresponding with our @ prior? classification and that no
single character is outstanding in accounting for the variation in principle
components one and two. Variation in principal component one is prima-
rily due to SCALL, INFLW, BCRAT, BRACTW, RAYL, BRACNL, CIRAT,
ACHNL, ACHNLW, SPIKLTW, PPEDN, and ASRAT, and variation in
principle component two is primarily accounted for by CULML and CULMW.
The results of DA are used to rank individual characters (Table 3) in order of
taxonomic reliability and to determine which combinations of characters
(Table 4) best classify our specimens into their categories as determined by PCA.

1052 Stipa 18(4

~

Taste 2. Eigenvectors and cumulative variance for first six principal components in principal compo-

nents analysis

Eigenvectors

Variable PCl PC2 PC3 PC4 PC5 PC6
PPEDN -0.245 -0.165 -O.155 -0.06] -0).232 -0.070
BRACNL -0.254 0.102 -0.065 0.001 0.140 0.147
CULML 0.167 -0.485 0.097 -0.039 0.275 0.007
BRACTL -0.226 -0.270 -0.134 -0.08 | -0.014 0.049
RAYL -0.255 -0.244 0.061 -0.042 -0.246 -0.077
INFLW -0.269 sOc 117 0.170 -O.017 -0.237 -0.060
SPKW -0.192 -O.1 1] 0.354 -0.293 -0.402 -0.100
CULMW -O.197 -O.410 0.005 0.224 0.113 -0.029
BRACTW -0,258 -O.181 -0.077 0.158 0.082 -0.090
LEAEW -0.196 -0.355 -0.060 0.230 0.254 -0.054
SPKLTW -0.248 0.172 O.151 -O.017 0.296 0.168
SCALI -0.274 0.182 0.149 0.041 0.150 47
ACHNI -0.252 O.131 -0,029 0.212 0.091 0.342
ACH NW 0.075 0.026 O.415 0.700 -0.241 0.272
ANTHL -0.137 0.256 0.054 0.316 0.161 -O0.825
SCALN 0.007 -0.060 0.676 -0.297 0.245 -0.03

CIRAT 0.253 -0.190 0.033 0.006 0.318 0.016
ASRAT 0.242 -0.187 -0.212 0.110 -().187 0.042
ACHLW -0.249 0.087 -0.160 -O0.181 0.278 0.102
BCRAT -0.262 0.123 -0.165 -0.049 -O.1 10 0.071

Cumulative
variance 0.544 0.654 0.728 0.785 O.831 0.867

DISCUSSION

Kiikenthal (1935-1936) treated Cyperms drummondii as a variety of C. virens
with no justification other than brief diagnoses of the taxa. Denton
(1978), in a numerical taxonomic analysis of the Lvza/ae group of Cyperis,
came to the same conclusion as Kiikenthal (1935-1936). Denton (1978, p.
257) stated that C. drwmmondii and other varieties of C. virens could not be
elevated in rank because of overlap in scale and achene dimensions in some
collections. However, Denton’s research involved little fieldwork and in-

cluded few C. drammondii specimens from the southeastern United States.

Our insite! included numerous recent collections of Cyperus drummondii
from the southeastern United States, not seen by Denton (1978). Discrete
clusters obtained with PCA indicate our specimens can be readily identified
as either C. drammondii or C. virens based upon our character set and that C.
drummondii and C. virens are distinct species. Further, DA shows that six
characters (SCALL, SPKLTW, ACHNL, ASRAT, CIRAT, BCRAT) are par-

ticularly effective in correctly classifying our specimens and that these six

1053

CARTER ET AL., Taxonomy of Cyperus virens and drummondii

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First Component
Fic. 1. Results of principle components analysis of Cyperus virens (Oj les) and C. drammondii

(closed circles).

liscrimt t lysis by individual characters.

Taste 3. Proportion of specimens correctly classified in

Proportion Proportion

Individual Correctly Individual Correctly

Characters Classified Characters Classified
SCALL 0.986 BRACNL 0.832
SPKLTW 0.968 PPEDN 0.828
ACHNL 0.953 RAYL 0.780
AT 0.946 BRACTL 0.752
BCRAT 0.915 ANT 0.726
IRAT 0.905 SP 0.720
ACHLW 0.885 LE 0.695
CULML 0.875 CULMW 0.649
BRACTW 0.849 ACHNW 0.615
INFLW 0.833 SCALN 0.548

characters are the minimum combination required to classify all of our specimens
into the two groups defined by PCA. Our specimens were classified with
99.3% accuracy using two combinations of four characters each (SCALL,

SPKLTW, ACHNL, ASRAT and SPKLTW, CIRAT, ASRAT, BCRAT), two
combinations of three characters (SCALL, SPKLTW, ACHNL and ASRAT,
BCRAT, CIRAT), and three combinations of two characters (SCALL, ASRAT;
SCALL, CIRAT; and SCALL, BCRAT). Moreover, DA showed the combi-
nation of key characters (SCALL, SPKLTW, PPEDN, RAYL, LEAF W, ASRAT)
used by Denton (1978) correctly classifying 99.1% of our specimens. Ad-

1054 Stipa 18(4)

TABLE 4. Proportion of specimens correctly classified in discriminant analysis by selected groups of

characters

Proportion
Grouped Correctly
Characters Classified

SCALL, SPKLTW, ACHNL, ASRAT, BCRAT, CIRAT, ACHLW, BRACTW,

INFLW, BRACNL, PPEDN, RAYL 1.000
SCALL, SPKLTW, ACHNL, ASRAT, CIRAT, BCRAT 1.000
SCALL, ACHNL, BCRAT, CIRAT, taal INFLW, BRACNL, RAYL 0.993
SCALL, SPKLTW, ACHNL, ASRA 0.993
SPKLTW, CIRAT, ASRAT, re 0.993
SCALL, SPKLTW, ACHNL 0.993
ASRAT, BCRAT, CIRAT 0.993
SPKLTW, ASRAT 0.993
SCALL, ASRAT 0.993
SCALL, CIRAT 0.993
SCALL, BCRAT 0.993
SCALL, SPKLTW, ASRAT, C ae BCRAT 0.986
SCALL, SPKLTW, ACHNL, AT 0.986
SCALL, SPKLTW, ACHNL, 0.986
BCRAT, CIRAT, BRACTW, me LW, BRACNL, RAYL 0.981

SPKLTW, CIRAT 0, ae

SPKLTW, BCRAT ).980

ditionally, the qualitative character, scale shape, was observed to be mark-
edly and consistently different between the two taxa and is included in the
dichotomous key.

In summary, of the six key characters used by Denton (1978), we found
three (SCALL, SPKLTW, ASRAT) to be highly reliable and concur with
their relative placement in Denton’s dichotomous key. Because of substan-
tial overlap between the two taxa, Denton’s remaining three key characters
(PPEDN, RAYL, LEAFW) proved to be of limited use in our study. Addi-
tionally, we have identified four other taxonomically useful characters: ACHNL,
BCRAT, CIRAT, and floral scale outline.

CONCLUSIONS

Results of PCA and DA show six characters are taxonomically useful in
separating Cyperus drummondii and C. virens. These in ranked order as deter-
mined by PCA and DA are SCALL, SPKLTW, ACHNL, ASRAT, BCRAT,
and CIRAT. Additionally, we have determined that the qualitative charac-
ter, scale outline, is useful in separating the two taxa. Further, the discrete
clusters obtained in PCA indicate C. virens and C. drummondii are distinct
in the southeastern United States and support recognition of C. drammondii

on)

at the rank of species as follows.

CARTER ET AL., Taxonomy of Cyperus virens and drummondit 1055
TAXONOMIC TREATMENT
Key to Cyperus virens and Cyperus drummondi
1. Floral scales oblanceolate, at least 1.7 mm long; spikelets at least 2.25 mm
wide; achenes less than 0.7 times as long as subtending floral scale; achenes
1.2-1.5 mm long; longest primary inflorescence bract length at least 0.45

times as long as culm; culm less than 8 times longer than inflorescence width;
1. C. virens

OQ

primary peduncles 5—
_ Floral scales ovate, less than 1.7 mm long; spikelets less than 2.25 mm wide;
achenes at least 0.7 times as long as subtending floral scale; achenes 1.1—-1.2
mm long; longest primary inflorescence bract less than 0.45 times as long
s culm; culm (8—)10—33 times longer than inflorescence width; primary

2. C. drummondii

—

peduncles 3—5
1. Cyperus virens Michx., Fl. Bor.-Amer. 1:28. 1803. (Fig. 2). Type: “Caro-

lina,” Herb. A. Michaux (HOLOTYPE: P, microfiche!).

Cespitose perennial herb. Roots fibrous, reddish brown. Culms 16-100
cm high, 3-13 times longer than inflorescence width, 2.4—6.9 mm wide,
triquetrous, scabrid. Leaves basal, blades 3.5-13.2 mm wide. Primary
inflorescence bracts 4-8, longest 17-75 cm long, up to 0.3-1.5 times as
long as culm, 2.7-13.2 mm wide. Inflorescence usually diffuse, (4.2—)7—
18.4 cm wide, rays 5—10, longest 2.4-12 cm long. Spikes globose, 1.2—7.3
cm wide. Spikelets flattened, 1.7-3.2 mm wide. Floral scales 12-38, alist

chous, spreading to ascending, bicarinate, oblanceolate, (1.35—)1.9-2.4 mm
long, pale grayish-green, turning brown. Stamens 1—2, anthers 0.8— 3 mm
long. Style 3-branched, exserted, deciduous. Achene oblong-ellipsoidal, 2.7—
4.1 times as long as wide, (0.9—)1.2-1.5 x (.3—0.4 mm, trigonous, stip!-
tate, brown.

Distribution and habitat —Widely distributed in the New World: south-
eastern United States, Caribbean, Mexico, Central America, and South America
(Denton 1978; Kiikenthal 1935-1936). In the coastal plain of the south-
eastern United States, from Texas through Florida into North Carolina (Fig.
3). Common ina variety of disturbed, low, intermittently wet habitats, such
as ditches, stream bottoms, edges of ponds and lakes.

Phenology. —Flowering and fruiting April through December.

Specimens examined. U.S.A. ea poe Co.: Point Clear, 22 May
Kral 55724 (VSC); Battleship oo ay, 1.1 mi E Mobile city limit, 8 Aug 1989, Carter
8100 (ctb, VSC). Butler Co.: SE € ele 12 Jul 19 . Kral 85208 (VSC). ones Co.:
SW Jackson, | Jun 1972 ee (VSC). Crenshaw Co.: N Rutledge, 23 Jul 1995,
Kral aes (VSC). Dallas Co.: SE Selma, 4 Oct 1994, a a 302 (VSC). eecice Co.:
ca. 2.5 mi SW Cottonwood, 26 Sep 1994, A MacDonald 7812 tele: VSC). Lee Co.: Uchee
Creek, - te 1960, Wills s.n. (SWSL); Spring Villa, 25 Jul 1977, H. Milas sm. (VSC).

obile Co.: Bayou la Batre, 7 Jun 1971, Kra/ 53056 (VSC). Montgomery Co.: WSW

Montgomery, 10 Oct 1970, Kral 41573 (VSC). | 2ike Co.: 5 mi SE Troy, 12 te 1973,

Kral 51217 (VSC). Froripa. Alachua Co.: NW Gainesville, 29 Jul 1978, Dunn 43 (ctb),

—

—_
Cy

1056 Stipa 18(4)

=

wise
se

Pic. 2. Cyperus drummondiz: A. Inflorescence. B. Floral scale and achene. C. Spikelet. Drawn
from Carter 10738 (ctb). Cyperus virens: D. Inflorescence. E. Floral scale and achene. F. Spikelet.
Drawn trom Carter 6961 (ctb). G. Habit: C. virens (left) and C. drammondii (right).

CARTER ET AL., Taxonomy of Cyperus virens and drummondii 1057

Fic. 3. Discribution of Cyperus virens in the United States.

6 May 1979, Dunn 595 (ctb); Paynes Prairie, 19 May 1979, sie 15 (FSU). Calhoun Co.:
E Bountstown, 6 Jun 1956, Redfearn 2197 (FSU). Columbia Co.: N Providence, 21 May

1964, Godfrey 63797 (FSU). Escambia Co.: Pensacola, 24 Jul 974 4, Godfrey 73759 (FSU).
Franklin Co.: NW Apalachicola, 29 Jun 1956, Kral 2737 (PSU); St. Vincent Island, 29
Jan 1987, Auden 10290 (FSU). hes — 6.5 mi NW Ape . achicola, 12 May 1986, Anderson

940] (FSU). Hillsborough Co.: .S mi NE of bridge, 18 May 1974, Burch ee (FSU).
Jackson ae S Sneads, 23 Sep 7 Gal 756063 ae Lake Co.: E Bar 10 Apr
1962, Godfrey 61885a (FSU). Leon Co.: Lake Bradford, 23 Jul 1957, ee 55675

(FSU); Apalachicola National Forest, 25 Jun 1 993, Godfrey 84612 (VSC); Tallahassee, 22
Jun 1989, Godfrey 83318 (VSC). Liberty Co.: N Bristol, 30 Sep 1964, Godfrey 64661 (FSU).
Madison Co.: Greenville, 24 Jun 1956, Godfrey & Kral 54922 (FSU); 14 Jun 1991, A
84184 (VSC). Okaloosa Co.: Niceville, 12 Oct 1974, Godfrey 73974 (FSU). Polk Co.
Lake Weohyakapka, Nalcrest, 7 May 1980, E.M. Wheeler s.n. (ctb, VSC). Santa Rosa
Milton, 2 Sep 1976, Godfrey 75489 (FSU). Wakulla Co.: Newport, 23 Jul 1961, Ward
2737 (FSU). GeoraiA. Baker Co.: Newton, 28 Sep 1991, Carter 9379 (ctb, VSC). Ben
Hill Co.: Red Bluff, 19 Aug 1967, Faircloth 4640 (VSC). Brantley Co.: 0.8 mi W Nahunta,
4 Jul 1988, Carter 6961 (ctb, VSC). B aie n Co.: Fe. Stewart Military Reservation, 24 Jun

1992, Carter 9930 (VSC). Camden Co.: Kinlaw, 7 Jul 1995, Carter 12523 (WSC). Cook
Co.: 6 mi W Adel, 19 Jul 1965, Fir ae eee Decatur Co.: 2.7 mi N state line,
20 May 1967, Faircloth 4466 (VSC). Glynn Co.: Sterling, Hwy. GA 99, 23 May 1997,

Carter 14032 with Alexander mre 7.72 mi a os Pleasant, Hwy. US 341, 23 May 1997,
Carter 14037 with Alexander (VSC). Grady Co.: 4 mi N Cairo, 7 Aug 1987, Carter 6010
(ctb, VSC). Liberty Co.: Ft. Stewart Military Reservation, 16 Jul 1992, Carter 10177 (VSC).
Lowndes Co.: N Valdosta, 20 Jul 1988, Carter 7092 (ctb, VSC); 11 Jun 1989, Carter 7892
(VSC). McIntosh Co.: Butler Island, 1.4 mi SW Darien, Hwy. US 17, 23 May 1997, Carter

L058 SIDA 18(4)

14030 with Alexander (VSC). Seminole Co.: 0.4 mi E Iron City, 5 Aug 1989, Carter 8023
aie VSC). Thomas Co.: 3.1 mi NE Metcalf, 2 Aug 1965, Faircloth 2628 (VSC). Turner
2.6 mt N Cravey, 28 Jun 1993, Carter 10843 with Kral (VSC). Wayne Co.: Gardi, 23
May 1977, Gave 14054 with Alexander (VSC). Wilcox Co.: 8.8 mi. N Abbeville, Hwy.
US 129, 9 Aug 1987, Carter 6060 (ctb, VSC); 1.5 mi N ject. Hwys. US 280 and GA 215,
30 May 1998, D. Alexander s.n. (VSC). we ISIANA. Acadia Parish: 0.5—1.0 mi E jet.
Hwys. LA 124 and US 90 in Estherwood, 3 Sep 1992, Carter 10444 (VSC). Allen Parish:
ca. 8 mi N Elton, Thzeret 29557 (FSU). fener Parish: Platternville, ca. | mi. N jet
Hwys. LA 70 and 308, 19 Sep 1986, Bryson 5054 (ctb). Beauregard Parish: near DeQuincy,
13 Jul 1963, Brown 17793 (ctb). Calcasieu Parish: ca. 1.5 mi NNW Starks, 18 Jun 1968,
Thieret 29424 (FSU); Chloe, 3 Sep 1992, Carter 10517 (VSC). Cameron Parish: ca. 10 m
W Holly Beach: 14 Dec 1957, Reese G Harris 1536 (FSU). East Baton Rouge Parish: E
Baton Rouge, 17 Nov 1993, McKenzre 1533 (VSC); Burden Research Center, S Hwy. 1-10,
28 Sep 1994, Bryson 14487 (ctb). Franklin Parish: SW Holly Grove, 17 Sep 1981, To-
mas 78546 (ctb). Jefferson Parish: 5 miS Marrero, 5 Jun 1948, Ewan 17710K (FSU). Lafayette
Parish: ca. 5 mi S Lafayette, 5 Oct 1957, Reese 1414 (FSU); 5 mi S Raynes, 15 May 1967,
Wooten 1042 (FSU); Lafayette, 16 Jun 1993, Carter 10739 (ctb, WSC). Livingston Parish:
1.3 mi N Bayou Manchac, 11 Sep 1979, Carter 2084 (WSC). Natchitoches Parish: Chest-
nut, Hwy. LA 9, 19 Sep 1981, Joye 2224 & Nielsen (ctb). Orleans Parish: New O: leans, 15
Oct 1993, Carter 11557 (VSC). Rapides Parish: Pineville, 16 May 1994, Bryson 13452
(ctb, VSC). Richland Parish: W Rayville, Hwy. US 80, 21 Aug 1990, Bryson 10203 &
Newton (ctb). St. Martin Parish: NW D ee Lake, 30 Oct 1963, Thieret 16611 (FSU); St.
Martinville, 16 Jun 1993, Carter 10751 (WSC). St. nae Parish: 2.7 mi N ats US
11 at Pearl ce 26 Aug 1991, Carter 8994 (ctb, VSC) Slidell, 3.1 mi. W jct
I- 2 and US 11, 17 May 1994, Bryson 13496 (ctb). oe Barish: 4 mi SW eee
2 May 1963, 160 (FSU); Lees Landing, 17 Sep 1993, Carter 11351 (VSC). Terrebonne
Parish: Brule, 7 Jul 1977, Webster 800 (ctb); Cocodrie, S Houma, 12 Aug 1989, Thomas
112004 (VSC). MISSISSIPPI. Bolivar Co.: S Shaw, 25 Jul 1994, Bryson 13982 with MacDonald
oS VSC). Clarke Co.: Pachuta, 28 Aug 1991, Carter 9087 & Bryson (ctb, VSC). Covington
».: Collins, 7 Dec 1994, Bryson 14705 (ctb, VSC). Forrest Co.: W Brooklyn, 4 fee 1992;
eile 11840 (ctb), Bryson 11852 (cth, ao Camp Shelby, NE McLaurin, Hartfield Creek,
15 Oct 1994, Bryson 14506 et al. (ctb),; Camp Shelby, NE Namib i. S of Forrest Ave, 15
Oct 1994, Bryson 14522 et al. (ctb). George Co.: 7.5 mi. ESE Lucedale, 19 Jul 1988, Morris
3224 (ctb). Hancock Co.: Buccaneer State Park, 26 Aug 1991, Carter 8976 & Bryson (ctb,
VSC); Hwy. MS 607, 1.4 mi ESE Hwy. US 90W, 26 Aug 1991, Carter ae (ctb, VSC);
Mississippi Welcome Center, S Hwy. I-10, W ee MS 6( : . Aug 1991, Carter 8988
(VSC), 17 May 1994, Bryson 13529 (ctb), W | Creek, 1 ct 1993, ane 13264 G
Carter (ctb, VSC). Harrison Co.: G ulfport, 6 es eo se : on (VSC), Bryson 11991
(ctb, VSC), Bryson 11997 (ctb, VSC), Bryson 11998 (ctb); Orange gs C eee Cen-
ter, 13 Oct 1993, Lockley sn. (ctb), 16 Oct 1993, Bryson 13165 (ctb, VS 7 Jul 1994,
Bryson 14004 (ctb); jot. Hwys. I-10 a MS 67, 27 a 991, Carter ben in tb, VSC).
Hinds Co.: Jackson, S of Lakeland Dr., 0.3 mi. W of Pearl River on Hwy. MS 25, 29 Oct
1993, nee 13300 Set ae Bl fo eas Park, 26 Aug 1995, Mac pees ge et. al,
(ctb). Humphreys Co.: 1.5 mi SW Isola, 23 Jun Bryson 8141 (ctb, VSC). Jackson
Co.: 7 mi E Moss Poine, 2: Jul 1989, ae 8666 (ctb, VSC); Pasc: aoe a, | . Sep 1991,
Bryson 11043 (ctb, VSC); 4 Aug 1992, Bryson 11879 (VSC), 9 Aug 1993, Bryson 12597
(ctb, VSC); Moss Point, | Sep 1993, Bryn 12841] (ctb, VSC); Be Point, 16 Sep 1993,
Carter 11316 (WSC). Jefferson Davis Co.: S Prentiss, 12 Aug 3, Bryson 12705 (ctb,
VSC). Lamar ie mi. N en Hwy. US 11, 18 May 1 a 4, Bryson 13568 (ctb).

_

CARTER ET AL., Taxonomy of Cyperus virens and drummondii 1059

Lawrence Co.: NE jct. Hwys. MS 27 and 44, 29 Jul 1994, ey 14094 es Hi Lin-

coln Co.: § Brookhaven, 29 Jul 1994, Bryson 14103 (ctb, VSC). Marion Co.: 4.6 mi. S jct.
Hwys. US 98 and MS 13, oo mi. S Upper Little River, S Sram 29 a 1994, Bryn
ida Newton Co.: 4 mi E Newton, 31 May 1985, Bryson 3943 (ctb, VSC). P

r Co.: 3 mi W Picayune, 37 Jun 1967, Sargent & Jones 13766 (FSU); aaa a0 ct
ee Bryson 13248 (ctb, VSC); 2.5 mi N Carriere, 26 Aug 1991, Carter 9005 (ctb, VSC).
Perry Co.: New Augusta, 19 Oct 1994, Bryson 14640 (ctb, VSC). Pike Co.: Percy Quinn
State Park, 9 Jun 1967, Jones 13229 (FSU), 28 Sep 1994, Bryson 14462 (ctb, VSC). Rankin
Co.: 0.5 mi § jet. Hwys. 49 and I-20, 22 Oct 1976, eee 790 (FSU); Flowood, 7 Sep

1993, Bryson 12973 (ctb, VSC); Richland, W Hwy. US 49, 1.2 mi. S jet. Hwys. I-20 vn
US 49, 27 Sep 1994, Bryson 14429 (ctb). Sharkey Co.: ca. , mi ENE Anguilla, 14 re
1988, Bryson 8348 (ctb, VSC); Delta National Forest, 3 Aug 1993, Bryson 12544 (ctb, VSC).
Smith Co.: SW Raleigh, 3.2 mi. W jct. Hwys. MS 35 and 540, 26 Jul 1995, Bryson 14952
(ctb). Stone Co.: Flint Creek Water Park, 18 Oct 1993, Bryson 13280 (ctb). Walthall Co.:
NW ES hashes 0. - mi. W jet. Hwys. US 98 — MS 583, 29 Jul 1994, Bryson 14089
(ctb). Washington . Leroy Percy State Park, 2-6 May 1975, Gunn 10637 (SWSL), 1
Jul me Ss nes 904 fo 16 Jul 1993, Bryson 12505 (ctb, VSC). Wayne Co.: Waynesboro,
| Sep 1993, Bryson 12934 (ctb, VSC). Wilkinson Co.: Little Buffalo Creek, N of Hiram
Mice Rd., 9 Jun 1991, Rosso 91-271 (ctb), Rosso 91-274 (ctb); ca. 8 mi N Woodville,
29 Sep 1994, Bryson fae cel VSC). NORTH CAROLINA. Carteret Co.: 3.6 mi NE
Beaufort, Hwy. US 70, 30 Jul 1988, Carter 7202 (VSC). Dare Co.: 2.6 mi W Alligator
Point, Hwy. US 64 oe 1988, Carter 7199 (ctb, VSC). Tyrrell Co.: 10.2 mi E Colum-
bia, ee US 64, 29 Jul 1988, Carter 7185 (ctb, VSC). SOUTH eee Barnwell
ane Barnwell, 12 Jun 1987, Horn 1904 (VSC). Berkeley Co.: Rte. 41, 5 ee Ree.

8 Jun 1983, ieNedas 35 (ctb). Colleton Co.: ca. 1 mi. "N Green "Pond, Re , 20
i 1993, MeNe#lus 93-1921 (ctb); vic. Green Pond, Rte 29 Aug 1994 a
94-857 (ctb). Dorchester Co.: SW Summerville, 17 May 1957, oe 26402 SU), Georgetown
Co.: Cat Island, 26 Jul 1990, Nefson 9392 (VSC). ee 4.6 mi NW Tillman, 9 Sep

1968, Leonard 1928 (VSC). TEXAS. Aransas Co.: Lamar, _ os 1992, pe 10620 (VSC).
Bastrop Co.: 1.8 airmiles N 37° W of RR trestle at Sayersville, 13 Jun 1986, Carr 7567
(VSC). Bee Co.: Naval Air Station Chase Field, 30 Sep 1991, Carr 11539 (VSC). Chero-

e Co.: 2.6 mi W Reklaw, 17 Aug 1989, Carter 8240 (VSC). Gonzales Co.: N Hwy ~US
I-10 and W wy. US 90, 11 ie ne Carter 8151 (ctb, VSC). Jefferson Cas Port Arthur,
10 Aug 1989, Carter 8140 (ctb, VSC). Matagorda Co.: NNE Palacios, Fm. Rd. 521, 13

Sep 1992, Carter 10632 (ctb, Montgomery Co.: ca. 9 mi. S Conroe, 28 oe 1988,
Hatch 5449 (ctb). ae Co.: along Hwy. I-10, milemarker 868, 19 Jun 1987, Levens
2616 (ctb). Polk Co.: 7.7 mi. S on Hwy. 59 from jet. with Hwy. 190 in Livingston, 11
May 1988, Jones 1494 é Wipff (ctb).

Zi a la Sete oe a Torr. & Hook. in Hook., Ann. Lyceum Nat. Hist.
York 3:437 (and oo 1836. Cyperus virens subsp. drummondi (Torr. &
aie Royanie: ae ):254. 1970. Cyperms virens var. Puma Tort. & Hook.)
Kiik., PAlanzenreich IV. eae - 181. 1936. Type: U.S.A. TEXAS: Drummond
449 (LECTOTYPE, typified by Denton (1978): K; ISOLECTOTYPES: B, GH, —
XF!).

os

ales robustus Kunth, Enum. Pl. 2: 41. 1837. Cyperus virens Michx. var. robustus (Xunth)
flanzenreich IV. 20 (Heft 101): 181. 1936. Type: BRAZIL: Brasilia, Se//o 5.x.
a HOLOTYPE, jide Denton 1978).

1060 Stipa 18(4)

STS GHEal
= >t
Zot: Eee = s

Fic. 4. Distribution of Cyperus drammondii in the United States.

Cespitose perennial herb. Roots fibrous, reddish brown. Culms 35—170
cm high, (8—)10—35 times longer than inflorescence width, 2—4.7 mm wide,
triquetrous, scabrid. Leaves basal, blades 2.4—11.2 mm wide. Primary
inflorescence bracts 3—5, longest 11.3—47.9 cm, up to 0.523 times as long
as culm, 1.9-6.7 mm wide. Inflorescence usually tight, 2.1—-8(-11.9) cm
wide, rays 3—5, longest 0.9-7.7 cm long. Spikes globose, 1.1—7 cm wide.
Spikelets flatcened, 1.5—2.2 mm wide. Floral scales 1 8—42, distichous, spreading
to ascending, bicarinate, ovate, 1.4—1.6 mm long, pale grayish-green, turning
brown. Stamens 1—2, anthers 0.8—1.2 mm long. Style 3-branched, exserted,
deciduous. Achene trigonous, oblong-ellipsoidal, 2.4—3.9 times as long as
wide, 1.0—1.2 x 0.3-0.5 mm, stipitate, brown. Fig. 2.

Distribution and habitat —Southern Georgia and northern Florida west-
ward through southern Mississippi and southern Louisiana into southeast-
ern Texas (Fig. 4). Also, known from Nicaragua, Jamaica, Surinam, Ecua-
dor (Galapagos Islands), and Brazil (Kiikenthal 1935-1 936; Denton 1978).
In the southeastern United States, found in the lower coastal plain along
t

_—

ne margins of flatwoods ponds, seepage slopes, and coastal prairies.
Phenology. —Flowering and fruiting May through October.
Because of its greater height and tighter inflorescence Cyperus drummondii
has a strikingly different habit from C. virens; this is best observed in the
field. Cyperus drummondii does not appear to be as weedy as C. virens. Cyperus

CARTER ET AL., Taxonomy of Cyperus virens and drummondii 1061

virens 1s much more common and widely distributed in the southeastern
United States, occurring in a variety of low wet disturbed habitats, espe-
cially ditches. In contrast, C. drammondii is less common and tends more to
be restricted to less disturbed systems such as margins of flatwoods ponds,
seepage slopes, and coastal prairies. Also, as summarized in the key, C. drwmmondii
differs from C. virens in its narrower spikelets, greater achene length to scale
length ratio, ovate scales, and fewer primary inflorescence bracts.

Specimens examined. U.S.A. ALABAMA. Baldwin Co.: Gulf Shores, S$ Intercoastal
Waterway, 3 Aug 1996, Kral 86381 (VSC); Gulf Shores, N Intercoastal Waterway, 4 Aug
1996, Kral 80382 (VSC). FLORIDA. Calhoun Co.: ca. Ocheesee Landing, 10 Jul 1976,
Godfrey 75220 (FSU). Escambia Co.: near Pensacola, Hwy. I-10, E of Hwy. FL 291, 8 Aug
1989, Carter 8085 (ctb, VSC). Franklin Co.: 3 airmiles N Apalachicola, 19 Jun 1986,
Anderson 9577 (FSU). Gadsden Co.: between Quincy and Gretna, 3 Nov 1965, Godfrey
67556 (FSU). Jackson Co.: 4.9 mi N Sneads ject. Hwys. US 90 and FL a 19 Sep 1988,
Carter 7627 (ctb, VSC); Lake Seminole, 6 Aug 1977, H. McIntyre s.n. (VSC). Suwannee
Co.: E Live Oak, 19 Aug 1976, Godfrey 75398 (FSU). GEORGIA. sae Co.: W Dixie,
old Hwy. US 84, 22 Jul 1995, Carter 12532 (ctb, VSC). oe Co.: Hwy. US 27, 4.3 mi
S Ausmac, 5 Aug 1989, Carter 8019 (ctb, GA, MO, SWSL, VDB, VSC); E Bainbridge,
Hwy. US 84, 13 Oct 1992, Kral 80044 (VSC). Dougherty Co.: 6 mi S Albany, Thorne
5002 (US); ca. 4 mi SE Putney, 18 Aug 1989, Norris & Owsley 5854 (personal herbarium
R.A. Norris). Pierce Co.: 3.4 mi NE jct. Hwys. GA 15-121 and US 84 in Blackshear, 19
Jun 1992, Carter 9861 (ctb, VSC); Hwy. US 84, 1.4 mi N Blackshear jct. with Hwy. GA
15-121, 19 Jun 1992, Carter 9863 (ctb, VSC); SW Blackshear, 2.67 mi NE Hwy. US 84
Satilla River br idge, 17 May 1997, Carter 14024 (VSC); Carter 14055 w eb red (VSC).
Tift Co.: 1.6 mi W Tyty, Hwy. US 82, 26 Jun 1993, Kral 82744 with Carter (VSC); Carter
10863 vith Kral (ctb, VSC). LOUISIANA. Acadia Parish: 0.5—1.0 mi E jet. Hwys. LA
1124 and US 90 in Estherwood, 3 Sep 1992, Carter 10455 (ctb, ae 0. a mi W Mid-
land, 17 Jun 1993, Carter 10761 (ctb, VSC); 5 mi. S Eunice and 5 mi. NE Iota near as
16 May 1994, Bryson 13478, Allen, Cascio & Geter (ctb, VSC). oe Davis Parish: 2
i N jct. Hwys. US 165 and LA 101 in Woodlawn, 7 Sep 1992, Carter 10533 (VSC), :
jee 1.0 mi W ject. Hwys. US 90 and LA 1126, 8 Sep 1992, Carter 10542 ( (VSC).
Lafayette Parish: between Duson and Scott, 0.5 mi E Has Hwys. US 90 and LA 724, 6
Sep 1992, Carter 10508 (ctb, WSC); Lafayette, Moore Park, 16 Jun 1993, Carter 10738
(ctb, VSC). Vermilion Parish: Abbeville, A.B. Langlois 85 5c (US); ca. 7.5 mi SW Gueydan,
8 Sep 1992, Carter 10546 with McInnis & Smith (VSC). MISSISSIPPI. Jackson Co.: Moss
Point, Hwy. MS 63 between Frederick St. and Martin Luther King Dr., 0.5 mi. S Escatawpa
River bridge, 16 Sep 1993, Carter 11314 (VSC), 11315 (ctb, SWSL, VSC); 16 Oct 1994,
Bryson 14544 & MacDonald (ctb, SWSL, VSC). Stone Co.: Hwy. 49 at 2.7 mi S jet. with
Hwy. 26 in Wiggins, 20 Aug 1999, MacDonald 13422 with Warren 2 Fe On OUTH
a Charleston Co.: N McClellanville, Godfrey & Tryon 724 (US). TEXAS. Goliad
Co.: 8 mi S Coleto Creek, § Victoria, Hwy. 77, Jones 719 (US); Aon = 5.5 mi W
Orange, i 50890 (US).

ACKNOWLEDGMENTS

Portions of this article were submitted by D.K. Alexander in fulfillment
of course requirements for Directed Study (BIO 495) at Valdosta State University

1062 Stipa 18(4)

and were presented in poster format at the 1998 Symposium for Under-
graduate Research sponsored by the Valdosta State University Council on
Undergraduate Research. Appreciation is extended to Ken Ledford for as-
sistance with data collection and Drs. S.D. Jones and G.C. Tucker for con-
structive reviews of the manuscript.

on

REFERENCES

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americana, Vol. 6: Alismataceae a Cyperaceae. Universidad Nacional Autonoma de México,
Cuidad Universitaria, Pp. 423-440.

ANONYMOUS. 1996. Minitab reference manual: release 11 for Windows!™M, Minitab Inc.,
State College,

Bryson, C.T. and R. Carrer. 1994. Nores on Carex, Cyperus, and Ky/finga (Cyperaceae) in
Mississippi with records of eight species previously unreported from the state. Sida 16:
171-182

Carter, R. 1990, (Vert (Cyperaceae), an overlooked species in cemperate North
merica. es 14 - 69- 77.
CHapMAN, A.W. 1889. Flora of the southern United States, 2nd ed. Ivison, Blakeman, and
Company, NY.
CHAPMAN, A.W. 1897. Flora of the southern United States, 3rd ed. Cambridge Botanical

Sup} sr Company. Cambridge, MA

Ciewett, A.E. 1985. Guide to the vascular plants of the Florida panhandle. University Presses
of Florida, Tallahassee

Corre, D.S. and H.B. Corrett. 1975. Aquatic and wetland plants of the southwestern

Uniced States. Stantord University Press, Stanford, CA.
Corrett, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular p

ants of Texas. Texas
eee aie Renner, TX.

Denton, M.F. 1978. 7 pee treatment of the Luvzua/ae group of Cyperus. Contr. Univ.
Michigan Herb. 11:197—

Goberey, R.K. and - eta 1979. Aquatic and wetland plants of southeastern United
States: Monocotyledons. University of Georgia Press, Athens.

Harcn, $.L., K.N. Ganput, and L.E. Brown. 1990. Checklist of the vascular plants of Texas.

MP-1655. Texas a cueae ee bac en Station, College Station

Jones, S.D., J.K. Wiper, and P.M. Montrcomery. 1997. Vascular pluses of Texas: A com-
prehensive checklist a. synonymy, bibliography, and index. University of Texas
Press, Austin.

Koyama, T. 1970. New combinations in the Cyperaceae of the Galapagos Islands. Madrofio
20:254.

KUKENTHAL, 1935— “ 36. Cyperus. In: A. Engler and L. Diel, eds. Das Pflanzenreich IV.
20 (Hett ic ): 1-67

McGivney, M.V. o A revision of the subgenus Eveyperas found in the United States.
Cath. Univ. Amer. Biol. Ser. 26:1—74.

Monr, C. 1901. Plant life of Alabama. Contr. U.S. Natl. Herb. 6:1—-921.

Seper, G.D.F. 1984. Multivariate oo John Wiley & Sons. New York.

SMALL, J.K. 1933. Manual of the pelea flora. New York

SngatuH, PHA. and R.R. SOKAL. 1973. Numerical Taxonomy. W. H. Freeman and Com-
pany. San Francisco.

CARTER ET AL., Taxonomy of Cyperus virens and drummondii 1063
Tuomas, R.D. and C.M. ALLEN. 1993. Atlas of the vascular fora of Louisiana, Vol. I: the
erns & fern allies, conifers, & monocotyledons. Moran Colorgraphic Printing, Baton

Rouge, LA.

Torrey, J. 1836. Monograph of North American Cyperaceae. Ann. Lyceum Nat. Hist. New
York 3:1-4

TUCKER, G.
Monogr. 43:1-213.

WUNDERLIN, R.P. 1982. Guide to the vascular plants of central Florida. University Presses

8.
C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot.

of Florida, Tallahassee.
WUNDERLIN, R.P. 1998. Guide to che vascular plants of Florida. University Press of Florida,

Gainesville.

Ww

1064 Sipa 1L8(4

BOOK NOTICE

Pritip W. Henprick. 1984. Population Biology, The evolution and ecology
of populations. 1998 printing. (ISBN 0-867 20-043-X, pbk.) Jones
and Bartlett Publishers International, 7 Melrose Terrace, London W6

7RL, ENGLAND and Boston, MA. $47.50. 445 pp., numerous figures.

This text isa comprehensive resource for the study of population genetics and population
ion Now in its 15‘ year, it has been reorganized into three parts that explore I) Basic
principles of genetics, Il) Population biology, and IH) The interrelationship between these
two fields. In each chapter you will find theoretical explanations, case studies, examples for
relevant models, and graphics selected to highlight important topics. Short problem sets
at the end of each chapter provide students the opportunity to practice using the models
presented in the text. Part one provides students with a solid foundation in the field of
genetics, with discussions covering major topics such as chromosomal variation, the Hardy-
Weinberg Principle, and gene flow. The section on Population Biology begins with a dis-
cussion of factors influencing species distribution and abundance, and of population den-
sity and dispersion before entering into a discussion of demography. This provides a useful
context for understanding growth models and population dynamics. A thorough presenta-
tion of interspecific competition, predator-prey interactions, and the theoretical models

used to describe these processes concludes part two. Part three builds on lessons from the
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cal genetics, and applied population biology. The text illustrates the combined use of es-
tablished theories and models with modern technologies to solve current problems in 1 ecol-
etek

a

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ogy ora a biology. It is a useful resource for students of genetics, ecology,
ecology alik

Contents for Population Biology, The et olution a of aaa: Preface, Introduc-
c Frequencies, Selection, Mutation, Gene Flow,
Traits, genes "Affecting Distribution

tion, Genetic Variation, Allelic and Genoty
Genetic Drift, Nonrandom Mating, ou ee
and Abundance, Population dae and Dispersion, Population Growth, Demogt raphy,
Interspecific Competition, Predator-Prey Interactions, Ecological Models of Selection, Adaptation
and Speciation, Molecular Variation a Delian The Evolution of Social Behavior, eines
Genetics and Life-History Evolution, Coevolution and the Evolution of Interspecific Inte
actions, Applied Population Biology, Conclusions to Part 3, Glossary, Bibliography, ve
swers to Numerical Questions, Author Index, and Subject Index.—Charlotte Bryant.

Stipa 18(4): L064. 1999

STRIKING SEXUAL DIMORPHISM
IN LINDERA SUBCORIACEA (LAURACEAE)

LORAN C. ANDERSON
Department of Biological Science
Florida State University
Tallahassee, FL 32306-1100, U.S.A.

ABSTRACT

Male and female flower morphology is documented for Florida plants, augmenting the
original floral analysis of this rare, dioecious species (Lindera subcoriacea). Sexual dimor-
phism is sia apparent; leaves on male plants are much larger than those on female plants.
Leaf surfaces of the two sexes also contrast strongly in features of trichome size, epicuticu-
evelopment of peristomatal rims. Such dimorphism

jon

lar wax serena and pattern), and c
has not been previously reported for the Lauraceae.
RESUMEN

Se documenta la Eonontoloei = flores masculinas y femeninas de Lindera subcoriacea en
plantas de Floric andlisis floral de esta especie rara y dioica. El dimorfismo

sexual es muy 4 specie las hojas de las plantas masculinas son mucho mayores que las de
las plantas femeninas. Las superficies foliares de los dos sexos también contrastan fuerteme

en cuanto a las caracterfsticas del tamafio de los tricomas, ceras epicuticulares Ce
y patrén), y desarrollo de los bordes peristomaticos. Este dimorfismo no habfa sido citado
previamente en las Lauraceae.

INTRODUCTION

Wofford described Lindera subcoriacea, the bog spice bush, in 1983. This
species is now known to occur in at least eight states, ranging in coastal
states from Louisiana to Florida north to Virginia (Bridges & Orzell 1989;
Bryson et al. 1988; McCartney et al. 1989; Sorrie 1993). Although the range
appears extensive, the plant is nowhere common. Most occurrences consist
of one or a few plants. McCartney et al. (1989) signaled its possible pres-
ence in Florida by listing some peninsular populations as “Lindera affin. swbcoriacea.”
Sorrie (1993) stated they have proven to be L. benzoin (L.) Blume, but these
problematic peninsular plants need further study. Unequivocal L. swbcoriacea
was reported by Anderson (1995) from a population in the Florida panhandle

More attention to the Florida panhandle plants is warranted because Wunderlin
(1998) did not list the species for Florida, and additional field surveys since
the initial report (Anderson 1995) have revealed new dimensions in the species’
morphology.

Sipa 18(4): 1065-1070. 1999

1066 Stipa 18(4)

MATERIALS AND METHODS

The original plane upon which the Florida panhandle record was based
(Anderson 1995) was destroyed by brush removal under the C/i/fonza canopy
along Metts Creek in Okaloosa County shortly after the collection was made.
Intensive survey along Metts Creek and nine similarly nearly pristine clearwater
streams in the general area (on Eglin Air Force Base) for additional bog spice
bushes yielded a count of 12 plants (six of each sex, restricted to Metts Creek).
James R. Burkhalcer collected a sample (originally identified only as Lindera)
from a presumably male shrub (sterile) on the campus of the University of
West Florida in Escambia County. After being alerted to its identity, he
searched the area thoroughly and reported (pers. comm.) that the plants
were no longer extant as a result of habitat alteration,

Flowering materials were preserved in FPA (5 pts formalin, 5 pts proprionic
acid, 90 pts 70% ethanol). Living plants were tagged at blooming time so
that they could be more easily located later in the season. Samples with mature
leaves were taken from only a few plants because of the species’ rareness.
Vouchers are at FSU unless otherwise noted. Leaf morphology (of the larg-
est leaf per specimen) was studied from me representing the geographical
range of the species. Leaf samples were cleared in NaOH and stained in safranin
for study of venation and trichomes. Other samples were sputter coated with
gold palladium, and micrographs were taken on a Joel JSM-840 scanning

on)

microscope operated at LOKYV.
RESULTS
Wofford (1983) reported the plants were up to 2 m tall (his original description
was based solely on specimens from Mississippi and Louisiana as this spe-
cies was not known to occur elsewhere at that time), whereas Sorrie (1993)
said that in North Carolina the plants grew up to 4 m tall and were multistemmed.
At Metts Creek 5-4

—

Florida) the blooming shrubs were 1.5—4.2 m tall with
only 2—4 stems per clump. These deciduous shrubs bloom in mid-March
(as they do in Mississippi) before leaf development.

Florida flowers of this dioecious species differ in several aspects from Wofford’s
description. Some of the differences in size may have arisen because his
measurements came from restored (boiled) flowers, whereas mine came from
preserved flowers. Wofford (1983) stated Hower buds were ca. 2.5 mm wide
and tepals were 2.2 by 1.8 mm. Further, in staminate flowers, stamens were
2.5 mm long, and the pistillodium was 1.2 mm long (Wofford 1983). In

—

our plants, staminate flowers were generally larger (2.8—3.2 mm long by
3. 8—4.5 mm wide at anthesis); outer tepals were 2.4—2.8 by 1.8—2.0 mm,
whereas inner tepals were slightly smaller (2.2—2.5 by 1.5-1.7 mm). The
nine stamens were 2.7—2.9 mm long; the innermost series of three had a

ANDERSON, Sexual dimorphism in Lindera subcoriacea 1067

pair of stalked (1 mm) = landular outgrowths. The glandular heads were
0.8 mm wide and had 3-4 irregular lobes.

The original dees (Woftord 1983) stated pistillate flowers were on
pedicels 1.5 mm long, with tepals slightly smaller that those of staminate
flowers, and stamens (staminodes) variously developed, often reduced to glands.
The style was 1 mm long, and the ovary elliptic and 1.0 by 0.6 mm. In our
plants, pistillate flowers were on ae 1.5—2.5 mm long. Flowers were

ea mm long ae e excluded) by 2.2—2 6 mm wide. The outer tepals
were 2.0—2.2 by 1.1—1.4 mm, and the inner tepals were 2.0 by 1.4—-1.5 mm
ee ne Dae ne outer tepals). The six outer staminodes (in two se-
ries) were reduced to naked filaments 1.4 mm long (no vestige of anthers),
whereas the three innermost staminodes (also filaments 1.4 mm long) had a
pair of flattened, stalked glands fused basally to the filament. The glands dif-
fered from those of the staminate flowers in that the heads (0.8 mm wide)
were broadly cordate and lacked irregular lobes. Styles were 1.4—1.8 mm long
and often curved. The elliptic ovaries were 1.6—1.8 mm long by 0.9-1.0 mm
wide. Mature fruits were scarlet, elliptical, and 10 mm by 6 mm

Wofford and Sorrie both gave the same range of 4—7.5 cm long by 2—3.5
cm wide for leaves of L. subcoriacea (Fig. 1). Leaves of female shrubs in the
Metts Creek population fall in that size range; the largest leaves per sample
(usually the penultimate on any given branch) measured 5.6—6.8 cm long by
2.4—2.8 cm wide (Figs. 2-4). This dioecious species exhibits sexual dimor-
phism, however; leaves on the male shrubs are 9.0—10.5 cm long by 3.5—4.6
cm wide (Figs. 5—7). The ranges in leaf size of the two sexes are almost com-
pletely nonoverlapping, if one discounts the very small broadly obovate leaves
that occur basally on the branches (they are 1-2 cm long on female shrubs
and 3.3—5 cm long on male shrubs). Incidentally, female plants of L. benzoin
var, pubescens from Florida have leaves larger than any seen in L. subcoriacea.

Mature leaves were dark green adaxially, whereas the abaxial (undersurface)
area was generally pale, grayish, glaucescent, and moderately appressed pubescent.
Venation is brochidodromous (Figs. |—7; Hickey 1979), and the areoles are
well developed. Trichomes were nonglandular and unicellular (Figs. 8—9),
as is typical for che Lauraceae (Metcalfe & Chalk 1979). Average trichome
length on leaves of four Florida female shrubs ranged from 0.19 to 0.22
mm long [and 0.23 on Wofford 82-121 (TENN), an isotype}, whereas tri-
chomes on Florida male shrubs averaged 0.29—0.32 mm long. The longer
trichomes on male shrubs also appear to be somewhat thicker than those on
female shrubs (Figs. 11—

Stained, partially cleared leaves showed the epidermal cells were thin-
walled; stomatal guard cells were 24—26 rm long, and the stomata were
paracytic (similar to those of fossil forms of Lindera illustrated by Bandulska

1068 Stipa 18(4)

7

Fic. 1-7. Comparative size (scale in cm) and shape of leaves (largest per branch) in Lindera
subcoriacea, Showing brochidodromous venation (i.e., secondary veins joined together in a
series of prominent arches). Fig. 1. Leaf from ae plant from George Co., MS, Wofford
82-121 (A, yore ve). Fig. 2-4. Leaves from female plants from Okaloosa Co., FL. Fig. 2
Anderson 14452. Fig. 3. Anderson 14920. Fig. 4. Anderson 15851, Fig. 5—6. ae from
male plants from Okaloosa Co., FL. Fig. 5. Anderson 14922. Fig. 6. Anderson 14921. Fig. 7.
Leaf from male plant from Escambia Co., FL, Burkhalter 5058.

926). The two subsidiary cells tended to be unequal in size on female leaves;
one cell was somewhat rectangular and lay parallel to the guard cell, whereas
the other subsidiary cell was more triangular in outline. Conversely, sto-
mata on male leaves had more evenly shaped, triangular subsidiary cells.

The grayish appearance of the leaf meer was due to wax accumu-
lacion more than to indumentum (Figs. 1O—16) as demonstrated by scan-
ning electron microscopy (SEM). The a layer of cuticular wax obscures
epidermal cell outlines. A papillose abaxial leaf surface is characteristic of
the Lauraceae (Metcalfe &Chalk 1979). The stomata on L. swbcoriacea leaves
are sunken and surrounded by prominent stomatal rims; peristomatal rims
are also evident—see Wilkinson (1979) for overview of these stomatal structures.
Stomatal rims and peristomatal rims may prove to be frequent among woody
taxa that have relatively firm (subcoriaceous) leaves in the southeastern United
States flora; elaborate rims have also been reported for Gordonia (Anderson
1983) and Sideroxylon (Anderson 1996).

Sexual dimorphism is also evident at the micromorphological level. Leaves
of female plants have extensive epicuticular wax papillae over the epider-
mal cells and the massive peristomatal rims (Figs. 10-12, 14-15). Leaves
of male plants have scattered epicuticular flecks of wax that do not hide the
cuticle, and peristomatal rims are not massive, but are present as 1—3 ridges
lying parallel co the long axis of the stomatal rim (Figs. 13, 16).

Unisexual flowers are, by definition, dimorphic, but in L. swbcoriacea the

ANDERSON, Sexual dimorphism in Lindera subcoriacea 1069

Fic. 8-16. Electron micrographs of leaf surfaces in Lindera subcoriacea. Figs. 8-9. Venation
and — x= 100. Fig. 10-13. Leaf surfaces showing cuticle and stomatal features,
» Fie Oe eat from female plant, Anderson 14452. Fig. 9. Leaf from male plant,
penne 14921 showing trichomes slightly longer and thicker than those on female leaf
(Fig. 8). Fig. 10. Female from George Co., MS, Wofford 82-121 (TENN, isotype). Fig
Female from Hoke Co., NC, Carter s.n. Figs. 12. Female from Okaloosa Co., FL, Anderson
14452, showing nearly continuous layer of epicuticular papillae and large peristomatal rims
(as in Fig. 10-11). Fig. 13. Male from Okaloosa Co., FL, Anderson 14921 showing scat-
tered epicuticular Hecks, reduced development of peristomatal rims, and thicker trichomes
(than female leaves, Fig. 10-12). Fig. 14. Leaf suraces of Anderson 14452 (female plant)
showing massive peristomatal rims covered with epicuticlar ae with stomatal rim
barely visible, x= 1500. Fig. 15. Portion of Fig. 14 showing detail of epicuticular papillae
and part of stomatal rim with the peristomatal rim, x = 3500. Fig. 16. Leaf surface of Anderson
149 31 (male plant) showing sparse eipcuticular flecks of wax and partial de ee of
peristomatal rim (forming brackets or ridges parallel to the stomatal rim), x= 1500

LO70 SIDA 18(4)

dimorphism extends to nonsexual parts of the flower with the different shapes
of glandular heads on staminodes. Sexual dimorphism in the Florida plants

is expressed vege cates in leaf size, trichome length, epicuticular wax patterns,
stomata (subsidiary cells), and peristomatal rim development. Vegetative
sexual dimorphism has been reported for L. benzoin (Cipollini & Whigham

1994), in which leaves on male shrubs averaged 1.3 times longer than those
on female shrubs. In L. swbcoriacea the dimorphism is more pronounced; male
leaves average 1.7 times longer than those on female shrubs. Apparently,
vegetative sexual dimorphism at the microscopic level has not been previ-

—

ously reported for Lindera or any other dioecious members of the Lauraceae
(Wood 1958; van der Werff 1991).
ACKNOWLEDGMENTS
My field work was sponsored in part by the Florida Natural Areas Inven-
tory through a grant from the U. S$. Department of Defense Legacy Pro-
gram. Kim Riddle and Bob Hursey assisted with electron microscopy, and
Ken Womble helped with graphics.

REFERENCES

ANDERSON, L.C. ite Trichomes and stomata of Gordonia lasianthus (Theaceae). Sida 10:103—113
5. Noteworthy plants from north Florida. Sida 16:581—587.
ron New geographical and morphological data for Sideroxylon thornei

(Sapotaceae). Sida 17:343-348
Bripoes, E.L. and $.L.ORZELL. 1989. Lindera subcoriacea (Lauraceae) new to Alabama. Phytologia

67:214—-216.
Bry SON, C.T., W. MCDEARMAN, and K. L. GORDON. 1988. Carex exilis Dewey (Cyperaceae)
a eee bogs. Sida 13:171-17
ae: M.L. and D.F. Wate HAM. ae {. Sexual dimorphism cost of reproduction in
the dioecious cae Lindera benzoin (Lauraceae). Amer. J. Bot. 81:65—75.
Hickry, L.J. 1979. A revised classification of the architecture of eee ae leaves. Pp.
25—39, In: C.R. Metcalfe and L. Chalk, eds. Anatomy of the dicotyledons, Vol. 1, 2nd

ed. Clarendon _ Oxford.
oe C.R. and L. CHatk. 1979. Anatomy of the dicotyledons, Vol. 1, 2nd ed. Clarendon
)xfor :
a ARTNEY, R.B., K. WourbaACcK, and J. Moore. 1989. Lindera in Florida. Palmetto 9(2):3—
SorRIE, BLA. 1993. Lindera subcoriacea Element Stewardship Abstract. The Nature Conser-
vancy, Sandhills Field Office, Southern Pines, North Carolina
VAN DER WerREE, H. L991. A to ous genera of the Lauraceae in the New World. Ann.

Missouri Bac Gard. 78:37
WILKINSON, H.P. 1979. The ne sae ice (mainly leaf). In: C.R. Metcalfe and L. Chalk,
nd

eds. Anatomy of the ne mei 5, Vol. 1, 2" ec / Cl. seen Press, Oxford. Pp. 97-165.
WOHORD, B.E. 1983. A new Lindera ( (Lauraceae) from North America. J. Arnold Arbor.
ih 2

33
Wooo, C E. 4 1958. The genera of the woody Ranales 1 in the southeastern United States.
J. Arnold Arbor. 39:296—346 (Lauraceae, pp. 326-3

NEW PLANTS FROM PAPUASIA. NOVELTIES
FROM THE LAKEKAMU AND
BISMARCK-RAMU EXPEDITIONS

W. TAKEUCHI
Botanical Research Institute of Texas
c/o Lae National Herbarium and
Papua New Guinea Forest Research Institute
P.O. Box 314, Lae, PAPUA NEW GUINEA
ABSTRACT

Two spray: are Ae aks descubed: rom the 1996 cee of the Lakekamu basin. The

new plants are and Trichospermum stevensii (Tiliaceae).
Medinilla bismarck-ramuUensis is ee ee from the 1995 biological survey of the Bismarck-
Ramu tract. Taxonomic and ecological notes are provided with the diagnoses.

ABSTRACT (MELANESIAN TOK PISIN)

| eae ]

is long ples Lakekamu, long Oktoba 14 igo Novemba

ee ce wile i ksmepimn ¢ tupela liklik diwai we i nupela samting olgeta. Lain
saientis givim ol nem olsem: Medinilla sapoi-riverensis na Trichospermum stevensii. Narapela
nupela samting kamapim a graun long Bismak-Ramu, na nem bilong dispela em i Medinilla

bismarck-vamuensts.
INTRODUCTION

The Lakekamu basin was botanically surveyed between October 14 and
November 12, 1996, as part of a general biological assessment organized
by Conservation International and the Foundation for People and Commu-
nity Development. The survey territory is regarded as one of Papua New
Guinea’s 16 terrestrial unknown areas (Sekhran & Miller 1995: 114) and is
also ranked among the highest value sites for biodiversity preservation in
Papuasia (ibid: 121). A synopsis of scientific results from the expedition
was previously published (Mack 1998). The present paper provides a for-
mal description for two of the plants listed as new species in the earlier publication
(Takeuchi & Kulang in Mack 1998).

A third species (Medinilla bismarck-ramuensis) is described from an ex
dition reviewed by Hedemark et al. (1997) and Takeuchi (1999).

De=

-_—

DESCRIPTION OF NEW SPECIES

MELASTOMATACEAE
Medinilla bismarck-ramuensis Takeuchi, sp. nov. (Fig. 1). Typr: PAPUA
NEW GUINEA. WesteRN HIGHLANDS Province: Bismarck Range, Mt. Oibo, ridgeline

Sipa 18(4): LO71-1LO81. 1999

1072 Stipa 18(4)

Fic. |. Medinilla bismarck-ramuensis Takeuchi. A. Vegetative habit. B. Leaf and petiolar
auricles. C. Inflorescence: flowers obscured by bracts. D. Cyme. E. Major stamen; perspec-
tives from the bud. Drawn from the type by N.H.S. Howcroft.

TAKEUCHI, New plants from Papuasia 1073
Camp 1, GPS 05° 35.513'S, 144° 47.252' E, elev. 2,357 m, 4 Oct 1995 (fl),
W. gees 10, 408 (HOLOTYPE: LAE; isotypes: A, K, L).

Medinillae schlechtero Mansfeld et M. mansfeldianae Merrill & Perry affinis sed abiea petiolis
auriculatis differt

Erect aside shrub to 2 m in height. Branchlets robust, terete, obliquely
ascending, indumentum setiform, spreading or subappressed, + dense; nodal
innovations crowded, setose, acroscopic, 10 mm long, glabrous and smooth
orat most obscurely plumulose near the base. Leaves isomorphic; blades fleshy
or coriaceous, elliptic or broadly lanceolate, 13.8-20.0 cm X 4.3—9.0 cm,
apex acute or gradually acuminate, base equal, cuneate to obtuse, margin
entire, not or hardly reflexed, lamina epunctate, bichromatic, adaxial sur-
face dark opaque green, abaxially pale purple-suffused, bifacially alveolate
in sicco; venation 5—7 plinerved, ventrally prominulous, more or less chan-
nelled, raised dorsally and distinctly purple, the lateral pairs of veins di-
verging from at most 22 mm above the base, at first subsericeous, progres-
sively setose towards the petiole, later glabrate on the mature lamina, reticulating
nerves invisible; petioles to 25 mm long, setose, proximally alate, the wings
paired, auriculiform, to 31 mm X 15 mm, foliaceous, the margins usually
entire, rarely erulose, surfaces purple and glabrous. Inflorescence dichasial, 2—
12 cm long, solitary or fascicled, variably inserted, infrafoliar and lateral to
cauline, occasionally from the axils of attached leaves, all parts dark red-
dish-purple except for the corolla, rachises cernuous, terete, not or only sparingly
branched, setulose at nodes, glabrous in between, conspicuously and per-
sistently bracteate; bracts geminate, orbicular, ca. 8 mm broad, involute,
infrequently with margins distantly fimbriate, surfaces glabrous, venulose,
nervation anastomosing; pedicels 3-4 mm, subtended by paired bracteoles.
Flowers (measurements from spirit-preserved material) acute in bud; hypanthium
setose, cupuliform-ovoid, typically 5 mm x 4-5 mm, margin denticulate
or irregularly notched; corolla 5-merous, choripetalous, white, imbricate,
petals 13 mm X 9 mm, asymmetric; stamens 10, glabrous, isomorphic but of
differing lengths, 5 long stamens alternating with 5 short ones, the longer outer
stamens: filament ca. 2.3 mm, anthers ca. 4.1 mm; shorter inner stamens:
filament ca. 1.9 mm, anthers ca. 3.0 mm; all stamens with anthers at first
inwardly retrorse and contiguous with the filament, dorsally ecalcarate or
nearly so, ventral appendages minute; style 7-8 mm long, glabrous, cylin-
drical, apically tapering, stigma discolorous and punctiform. Fruits not seen.

Distribution and ecolagy.—Medinilla bismarck-ramuensis is known only from
montane forests in Western Highlands and Morobe Provinces. The type was
found as scattered plants in fog-swept clearings, growing on mossy stumps
and logs.

Etymology.—The new species commemorates the 1995 Bismarck-Ramu
expedition.

1074 SipA 18(4)

Paratype. PAPUA NEW GUINEA. Morose Province: Aseki Patrol Area near cee
elev. 5,900 ft (1,800 m), 4 Apr 1966 (Al), L.A. Craven & R. Schodde 1157 (A, CANB, K, L,
LAE).

Medinilla is comparable to Cyrtandra in its pattern of endemic diversification.
Even at the time of the first revision of Papuasian taxa (Mansfeld 1925),
Medinilla was already one of the larger woody genera, with a conspectus of
53 species. Merrill and Perry (1943) later added 16 species using a key patterned
on Mansfeld. Baker (1916), Ohwi (1943), and Bakhuizen (1943) also con-
tributed species from the Wollaston expedition, the Kanehira-Hatusima
collections, and Dutch New Guinea, respectively. Numerous contemporary
gatherings cannot be accommodated by these earlier contributions, sug-
gesting that the number of undescribed species is substantial.

In Mansfeld (1925: 115-18) the new species will key to M. schlechteri
Mansfeld. Unlike most treatments from the ‘Beitrage zur Flora von Papuasien’
series, Mansfeld’s key is written in Latin rather than German. Although
outdated, Mansfeld’s treatment still stands as che most comprehensive available
account. Medinilla bismarck-ramuensis can be inserted into the couplet train
at species 50, in the following manner:

Folia supra glabra, subtus nervis pilis nonnullis vestitis
Petiolus auriculatus: Medinilla bismarck-ramuensis Takeuchi
Petiolus non auriculatus: Medinilla schlechteri Mansfeld

In Merrill and Perry (1943: 427-28), M. mansfeldiana Merrill & Perry is
the closest match to the new species, but the longer and manifestly auricled
petioles on M. bismarck-ramuensis are distinctive. The novelty’s affinity is
actually to a group consisting of four species (fork G, ibid: 42

Medinilla bismarck-ramuensis may be connected to the complex character-
ized by persistent bracts and bracteoles, and regarded by Merrill and Perry
(ibid: 422) as comprising a distinct section. In Papuasia, this alliance is
characteristic of the archipelagic stations from New Britain to the Solomons
and (unlike the new species) is usually markedly anisophyllous. It was not
previously thought to occur on the New Guinea mainland.

—

Medinilla sapoi-riverensis Takeuchi, sp. nov. (Fig. 2). Type: PAPUA NEW
GUINEA. GuLt PROVINCE: Lakekamu, near Sé a River, first ravine on the track above
Ivimka Research Station, 07° 44'S, 146° 30' E, elev. 183 m, 22 Oct 1996 (fl, fr),
Takeuchi & J. Kulang 11,398 (HOLOTYPE: ce ISOTYPES: A, ees BRIT, CANB,

Medinilla warica Mansteld similis sed inflorescentiis non terminalibus.

Epiphytic shrub, ascending or with branches extended and hanging. Stems
quadrangulate, tetrapterous, weak, hollow, surfaces green, minutely and distantly

stellate-furfuraceous on apical intervals, older parts glabrous and pustulate,
nodal innovations light-brown setose, 15-25 mm long, the youngest ones

Takeucui, New plants from Papuasia 1075

5mm

food!
Midige

y ee Ent T ea op

Fic. 2. Medinilla sapoi-riverensis Takeuchi. A. Veg
C. Mature infructescence. D. Flower, showing 3 petals, 2 stamens, and the style; other parts
removed for clarity. E. Fruit. Drawn from the type by N.H.S. Howcroft.

1076 SIDA 18(4)

somewhat plumulose but with the hairs fugacious. Leaves equal, obliquely
diverging; blades linear-e

oer

liptic or narrowly lanceolate, subcoriaceous, 23—
33cm X 5.2—8.0 cm, adaxial surfaces opaque green, abaxially pale green;
apex attenuate, prolonged into a filiform cauda or not, margins reflexed,
base cuneate to obtuse, lamina reddish-purple in flush, initially stellate-
pubescent on veins, glabrous when mature; venation 5-plinerved, the lat-
eral pairs diverging from less than 12 mm above the petiole, raised on both
sides, tertiary nervation transversal, scalariform, ventrally prominulous, dorsally
less apparent; petiole 8-15 mm long, glabrescent. Inflorescence ramigerous,
paniculiform, ultimately cymose, lax, pendulous, 10-18 cm long, ternate
or quaternate on the first order ramification, lateral branches to 4 cm long,
all axes nitid, red, flaccid, delicate, inconspicuously and stellately pubes-
cent, eventually glabrate; floral bracts linear, persisting, hardly visible; pedicels
1-3 mm long. F/owers (measurements from spirit-preserved material): hy-
panthium cylindric to narrowly ovoid, 3.0—3.5 mm x 2.5—2.8 mm, mar-

1

gin truncate or 4-denticulate, exterior surfaces light green, at first very sparsely
lepidote like the pedicel, glabrate at anthesis; corolla 4-parted, acute in bud,
petals pink, reflexed, ecostate, asymmetric, distally oblique, apiculate, 4.0
mm x 2.5—2.8 mm; stamens 8, equal, glabrous, filaments 2.3—2.6 mm long,
apically attenuate, anthers 1.9-2.0 mm long, distinctly calcarate and bi-
appendiculate, dorsal spur ca. 0.4 mm long, linear, basally directed, ven-
tral appendages lobulate, introrsely projecting, deltoid, ca. 0.2 mm long;
style 6.5—6.9 mm long, stigma discolorous and punctiform, receptive sur-
face obscurely papillate. Fruit globose, 5-6 mm in diameter, baccate,
polyspermous, exocarp green turning black when ripe; seeds oblongoid, 1.0—
1.2 mm X 0.5 mm, testa pale, verruculose.

Distribution and ecology.—Known with certainty only from the type lo-
cality. Medinilla sapoi-riverensis is infrequent in natural growth foothill for-
est, favoring mainly the wetter sites afforded by dark draws and gullies,
especially near flowing creeks with closed overstories. It is shade-loving and
does not occur in seral situations. A collection from Western Province (Henty
et al. in NGF 42788) may be M. sapoi-riverensis, extending its range further
west towards the border with Irian Jaya. The vegetative characters are con-
sistent with the new species, but che specimen lacks the fertile structures
required for reliable identification.

Etymology.—Medinilla sapoi-riverensis is named after the type locality in
the Lakekamu basin.

Medinilla sapoi-riverensis is distinguished by its elongate leaves and qua-
drangular-alate branchlets. Other salient characters are the lax, pendulous
inflorescences with verticellate branching. Innovations are setose at older
nodes and only caducously plumulose in the apical tufts.

Medinilla sapoi-riverensis keys to M. warica Mansfeld (Mansfeld 1925) but

TAKEUCHI, New plants from Papuasia 1077

the latter species has a terminal inflorescence. There is also an apparent affinity
to M. mansfeldiana Merrill & Perry, and M. schlechteri Mansfeld. The new
species differs from M. mansfeldiana by the nat 1 density of the indumentum,
length of the inflorescence, and size of the bracts. From M. schlechteri, it differs
in having clearly petiolate leaves, smaller bracts, and esetose calyces.

TILIACEAE

‘Trichospermum stevensii Takeuchi, sp. nov. (Fig. 3). Tyee: PAPUA NEW GUINEA.
ULF Sie Lakekamu, Sapoi River, streambanks near base camp, 07° 44'S, 146°
29.5'E, elev. 105 m, 7 Nov 1996 (fl bud, fr, xylarium collection), ia Takeuchi & J.
Kulang 11, oe (HOLOTYPE: LAE; tsorypes: A, BISH, BRIT, CANB, K, L).

1] ] ] pela
1 L

Small tree to 8 m tall. Stew without buttress or basal swell, exterior sur-
faces pale grayish-brown, unfissured, not lenticellate; bark thick, fibrous,
extractable in flexible strips or plates; wood white, weak, very light in weight.
Branchlets flexuous, terete, sparingly mucilaginous from cut surfaces, pithy,
indumentum stellate, squamulate, scales dark brown, at most 0.3 mm di-
ameter, coarsely-armed, crowded, occasionally accompanied by larger as-
cending hairs especially at the stem apices. S/puw/es caducous or persisting
through several nodes, oblong-ovate, typically 10 mm x 4 mm, rarely to
14 mm X 10 mm, foliaceous, yellow-green, cordately-based or auriculate,
proximally and laxly provided with indumentum like the branchlets, oth-
erwise glabrescent. Leaves distichous, membranaceous, adaxially dark dull
green, abaxially medium green and marked by opaque punctulations, ven-
tral surfaces puberulous, the hairs simple or stellate, following veins, dor-
sal surfaces stellate-pilosulous, hairs 4—8 armed, ascending, processes short,
ca. 0.2 mm long, the midrib also provided with a secondary indumentum
of smaller appressedly stellate scales; mature blades oblong, weakly con-
stricted near the middle, 22-32 cm x 10.5-14 cm; apex long-acuminate,
more or less abruptly developed, the acumen 2.0—3.5 cm long; leaf mar-
gins entire to the naked eye, remotely and minutely toothed under magnification,
serrulations glandular-thickened from excurrent nervules; leaf base trun-
cate or more commonly broadly cordate, the sinus when present to 1.5 cm
deep, basal glands marginal, occurring as linear callosities flanking the petiole
insertion; lateral veins 7—9 above a basal nervation, often in opposite pairs,
acutely diverging (ca. 45° from the midrib), generally straight, ending in
commissural looping nerves or not, major veins plane or hardly raised ven-
trally, dorsally prominulous and with primary axils stellately comose, higher
order axils clear, intercostals obliquely scalariform, tertiary venation lax;
petiole 14-25 mm long, unchannelled, indumentum as the branchlets. Inflorescence
axillary, to 8.5 cm length, axes texturally scabrid, variably covered by stel-
late scales and occasional interspersed hairs, rachides unequal, often fractiflex,

Folia subter consperce stellato-pilosa et costa mediz

1078 Sipa 18(4)

er | {oe
10 mm 1mm

Lh le axe off

3/9

Fic. 3. Trichospermum stevensii Takeuchi. A. Flowering branchlet. B. Mature leaf. C.
Vegetative branchlet and stipules. D. Infructescence. E. Dehisced fruits. F. Detail of cap-
sule from side. G. Capsule hairs, showing the 2 stellate types discussed in the text. Drawn
from the type by N.H.S. Howcroft.

TAKEUCHI, New plants from Papuasia 1079

bifurcate or ternate at the first branch, ultimately cymose, peduncle 5—30
mm long; bracts deciduous, free, linear-acuminate or ligulate, usually 2—3
mm long, sparsely stellate-hairy, the cymes subtended by 6 bracteoles. F/owers
not seen at anthesis, immature buds (all measurements from rehydrated material)
ovoid to subglobose, 4.5-5.5 mm x 5.5—6.0 mm, yellow-green; sepals 5,

valvate, fleshy, ovate, 5 mm x 3 mm, outer surfaces coarsely and minutely
stellate-hairy, inside non-stellately pilosulous with submoniliform hairs; corolla
included, imbricate, 5-choripetalous, chartaceous, reticulately venose, el-
liptic to oblong, ca. 4 mm x 2 mm, dorsally with indumentum like the
contiguous calycine surface, ventrally glabrate; androecium polyandrous,
congested, glabrous, anthers medifixed, arcuate; ovary compressed, pulverulent,
style 4-lobulate; receptacle pilose; pedicels ca. 1.5 mm long. Capsules 2-
celled, loculicidal, polyspermous, distinctly broader than long, 13-15 mm
x 20-23 mm, compressed but with otherwise weakly inflated lobes, sum-
mit retuse, style persisting, capsular base obscurely stipitate, androecium
residue present on the fruit, valves 2, thin-crustaceous, marginate, brown-
ish-green turning black after seed dispersal, persistently and basally con-
nate, inside glabrous, smooth or intermittently striate, outside somewhat
harshly tomentulose, the indumentum stellate, dimorphous, bilayered, the
larger hairs lax, 4-8 armed, processes 0.5—1.0 mm long, radiate, underneath
with a congested layer of coarse scales 0.3 mm diameter . Seeds obovate or
oblanceolate, dorsally convex and ventrally +cotyloid, ca. 1.4 mm xX 1.0
mim, testa brown, shaggy-fimbriate on equatorial margins, fimbriae fulvous,

3—4 mm long.

Distribution and ecology.—Trichospermum stevensii is thus far known only
from lowland rainforests of Gulf Province. The new species is characteristic
of regrowth environments such as gaps, landslides, forest margins, and surge
zones along open streambeds. It is most commonly seen in association with
various species of Macaranga, or as a co-dominant in early riverine succes-
sions. Nearly all sightings were in sterile condition during the survey.

Etymology.—The new species is named after Professor P.F Stevens, an authority
on the Malesian flora and a specialist in Ericaceae and Clusiaceae.

Paratype. PAPUA NEW GUINEA. Gulf ani baen subdistrict Baimuru, above Purari
River, 63 km at 65° from Baimuru, 07° 15! 45° 20' E, elev. 2: ‘ m, 26 Mar 1974 (fr,
wood), J.R. Croft et al. in LAE 61143 (A, PRE : ANB, K, L, LAE

In Kostermans’ (1972) revision of Trichospermum, T. stevensii arguably keys
closest to the extra-Papuasian 7! morotazensis Kosterm. and T! fosbergii Kosterm.
Although Kostermans’ reliance on underleaf indumentum is useful for pri-
mary separations, Trichospermum stevensti falls between the cracks on the first
order couplets. It is nearest to fork 1c (ibid: 405—406) but the combination
of dorsal scales with scattered, erect hairs is not covered by any of the prin-

i

1080 Sipa 18(4
cipal leads. The strictest solution for integration into the existing key is to
introduce another descriptor for the new species, as follows:

lf. Lower leaf surface squamulate on costae, and with scattered, erect stellate
hairs on intervenia and higher order nervation T. stevensii Takeuchi

If the underleaf hairs on the new species are generously interpreted, it
can be forced into Kostermans’ conspectus, in which case an apparent con-
nection to T) morotarensis and T: fosbergii will emerge. Substantive distinc-
tions in leaf and capsule size would still provide separation against the pre-
ceding taxa. Among Papuasian representatives, 1. sfevensi2 appears close to
T. tripyxis UX. Schum.) Kosterm., but the latter species has trivalved cap-
sules. The compressed bivalved fruits of T. stevensii also suggest affinity to
T. peekelv Burret, but the indumentum is clearly inconsistent with that species.
The type collection has branchlets with appressed stellate scales rather than
the erecto-patent vestiture on most Papuasian taxa.

The bast of Trichospermum stevensii is very resilient; the entire bark is stripped
in sheets from the trunk and the resulting mats applied as flooring by Lakekarmu
villagers. Other species of Trichospermum are noted for their fiber value (ibid:
403).

ACKNOWLEDGMENTS

Funding for my participation on the Lakekamu survey was provided by
Conservation International. Andrew Mack and the Foundation for People
and Community Development were the principal organizers for the survey.
The 1995 Bismarck-Ramu Expedition was funded by the Global Environ-
ment Facility and the United Nations Development Program. My work in
Papuasian botany was supported by the Liz Claiborne and Art Ortenberg
Foundation. Processing and identification of plant specimens occurred at
the Lae National Herbarium and PNG Forest Research Institute. Emily
Wood and Michael Bartley of Harvard Herbaria kindly responded to re-
quests for specimen information. Neville Howcroft provided illustrations
of the new species. John Pipoly HI, my colleague at the Botanical Research
Institute of Texas, corrected the Latin. My sincere thanks to all these indi-

viduals and organizations.

REFERENCES

Baker, E.G. 1916. Melastomataceae. In: Sate! = H. Report on the botany of the Wollaston
Expedition to Dutc ew Guinea, 1912 . Trans. Linn. Soc. London, Bot. 9:50—57
BAKHUIZEN VAN DEN ane R.C. 1 194. 43.A ete to the knowledge of the Ve eeee
occurring in the Malay Archipelago, especially in the Netherlands East Indies. Thesis
Utrecht (reprinted in Recueil Trav. Bot. Néerl. 40:1—391. 1946).

Hepemark, M., S. HAMILTON, and W. TAKEUCHI. 1997. Report on the firsc Bismarck-Ramu
biological survey with sociological and logistical comments. PNG Dept. of Environ-

ment and Conservation, Port Moresby.

TAKEUCHI, New plants from Papuasia 1081

KosTERMANS, A.J.G.H. 1972. A Sane of the Old World species of Trichospermum Blume
(Tiliaceae). Trans. Bot. Soc. Edinb. 41:401—430.

Mack, A. (ed.). 1998. A biological assessment of the Lakekamu Basin, Papua New Guinea.

Rapid Assessment Program Working Papers no. 9, Conservation International, Wash-
ington, D.C.

MansreLp, R. 1925. Di vacate cs von Papuasien. Bot. Jahrb. Syst. 60:105—148.

Merritt, E.D. and L.M. Perry. 1943. Plantae Papuanae Archboldianae, XII. J. Arnold
Arbor. 24:422-439

Onwi, J. 1943. The Kanehira Hatusima en ea ae of New Guinea plants. XVI.
Melastomataceae. Bot. Mag. (Tokyo) 57 (673

SEKHRAN, N. and S. MILLer (eds.) ne Papua — Guinea country study on biological
diversity. Dept. of Environment and Conservation and the Africa Centre for Resources
and Environment, reprinted 1996.

TAKEUCHI, W. 1999. Botanical al from the 1995 Bismarck-Ramu expedition in Papua

New Guinea. Sida 18:751-—7%

Takeucui, W. and J. KULANG. ee Vegetation Part 2: Botanical survey. In: Mack, A., ed.
A biological assessment of the Lakekamu Basin, Papua New Guinea. Rapid Acasa
Program oo apers no. 9, Conservation International, Washington, D.C. Pp. 36—
39, LO5—

1082 Stipa 18(4)
BOOK NOTICES
LORRAINE DASTON and KaTrHaRINeE Park. 1997. Wonders and the Order
of Nature, 1150-1750. (ISBN 0-942299-90-6, hbk.). Zone Books,
611 Broadway, Suite 608, New York, NY 10012. Distributed by MIT
Press, Cambridge, MA. $34.00. S11 pp., illustrated.

—

From dustjacket: “Wonders and the Order of Nature is about the ways in which European
naturalists from the High Middle Ages through the enlightenment used wonder and won-
ders, the passion and its objects, to envision themselves and the natural world. Monsters,

| | rt le
LIITCSE WCIC LIIC Tildal VOLS

gems ae it shone in the dark, petrifying springs, celestial apparitions
that adorned romances, ges philosophers, lured collectors, and frightened the devout.
Drawing on the histories of art, science, philosophy, and literature, Lorraine Daston and
Katharine Park explore and ex mi; ain how wonder and wonders fortified princely power, re-
wove the texture of scientific experience, and shaped the sensibility of intellectuals. This is

a history of the passions of inquiry, of how wonder sometimes inflamed, sometimes damp-
ened curiosity about nature’s best-kept secrets. Refracted through the prism of wonde rs,
the order of nature splinters into a spectrum of orders, a tour of possible worlds.

Contents for ee ~~ the Order of Nature, 1150-1750: Preface, Introduction: At the
Limit, The Topography of Wonder, The eae! of Things, Wonder Among the Phi-

losophers, Marvelous Particulars, Monsters: A Case Study, Strange Facts, Wonders of Art,
Wonders of Nature, The Passions of Inquiry, The I Shennan and the Anti-Marvelous,

Epilogue, Photo Credits, Notes, Bibliography, and Index.

Apbet JALILI and Zipa JAMzab. 1999. Red Data Book of Iran. A Prelimi-
nary Survey of Endemic, Rare & Endangered Plant Species in Iran.
With Contributions from S.C. Shaw, A.A. Maassoumi, U. Assri, N.
Mazhari and A. Rahmanpour. (ISBN 964-473-061-5, hbk.) Research
Institute of Forests and Rangelands, P.O. Box 13185-116, Tehran, IRAN.
Iran 85000 R; Outside $100.00. 748 pp., 85 mostly color drawings

and a few color photos.

Iran supports a total of around 8,000 plant species, and is one of the major centers of ende-
mism (1,727 endemic in Iran) in that pare of the world. The cotal number of listed taxa
based on defined IUCN categories, endemics, vulnerable, and endangered is 2,405. For
the category Vulnerable, 432 taxa are listed and 21 listed for Endangered. Contents for
Red Data Book of Iran. A Preliminary Survey of Endemic, Rare & Endangered Plant Spectes in
Iran: List of Figures, Introduction, The Phytogeographical Regions of Iran, Data Collec

tion and Analysis, Results and Discussion, Concluding Remarks, Species — eee
1, References, and Index. For each species, the t ee information is provided: Status
(Endangered, Vulnerable, Lower Risk, Data Deficient; Life Form, Distribution; vand Habitat.
The book is printed on quality paper and the color paintings sand photos are very nice.

Sipa 18(4): 1082. 1999

A NEW VARIETY OF PINUS GREGGII
(PINACEAE) IN MEXICO
JEFFREY K. DONAHUE
Boise Cascade Corporation
P.O. Box 1060
De Ridder, LA 70634, U.S.A

JAVIER LOPEZ UPTON
Especialidad Forestal, IRENAT

Colegio de Postgraduados en Ciencias Agricolas
Montecillo, Mex. 56230, MEXICO
ABSTRACT
new variety of Pinus gregeii is described. Variety australis is endemic to east central
Mexico and represents the southern disjunct region of the natural range of the species.

Variety gregei/ represents the northern population of the species. Characters that segregate
the varieties are presented
RESUMEN

Se describe una nueva variedad de Pinus ereggit. La variedad australis es endémica del
centro-este de México, y abarca la distribucién natural austral de la especie. La variedad
greggii representa la poblacién localizada en el norte de su distribucién natural. Se incluyen
caracteristicas que distinguen las dos variedades.

Pinus gvegeit Engelm. occurs in two disjunct regions in Mexico (Fig. 1)
separated by approximately 300 km, or four degrees latitude. Northern and
southern populations of P. gregg// grow in distinct environments. Northern
populations occur in the northern Mexico states of Coahuila and Nuevo
Leon, at elevations ranging from 1900 to 2600 meters above sea level. The
average annual temperature at the northern sites is 14° C, and they receive
an average annual precipitation of 650 mm (Donahue & Lopez Upton 1996).
Topsoils at northern P. greggiz sites are predominately neutral or slightly
alkaline (Donahue 1993).

Southern populations of P. gregg7/ occur in the central Mexico states of
Hidalgo, Puebla, Queretaro, San Luis Potosi and Veracruz at elevations ranging
from 1100 to 2400 meters above sea level. The average annual tempera-
ture at the southern sites is 17° C, and they receive an average annual pre-
cipitation of 800-1600 mm (Donahue & Lopez Upton 1996). Topsoils at
the southern P. gregg7z sites are predominately acidic (Donahue 1993).

A comprehensive study of the species began after differences in growth
rate, needle length and needle color became apparent in genetic trials planted

Stipa 18(4): 1083-1093. 1999

1084 Sipa L8(A)

N
km
— vy Peer
Y
s

Pinus greggii var. greggii

Pinus greggii var. australis

Gulf of Mexico

Pacific Ocean

162 N

110° W 102° 94°
re

rt A rN A

Fic. 1. Geographic distribution of Pius greggii in Mexico.

in Brazil, Chile, Colombia, South Africa (Dvorak et al. 1996) and Mexico
(Lopez Ayala 1998: Alba Landa et al. 1998), where trees from northern and
southern populations were planted side-by-side. The differences observed
in these field crials suggested that two distinct, unrelated taxa had been
planted by mistake. Studies of geographic variation in leaf, cone and seed
morphology, seed production and terpene chemistry were performed to determine
what differences existed, and to quantify them. Results from our compre-
hensive study indicated that only P. gregg/7 was included in the trials, but
several character differences exist between northern and southern popula-

tions suggesting recognition taxonomically

MATERIALS AND METHODS

Field sampling was done in six northern populations and six southern
populations from a wide range of geographic locations and elevations (Table
|). Leaf specimens were collected from 172 trees, and cones from 177 trees
of Pinus greggii. The samples were collected from the upper 1/3 of the crown
from healthy dominant and co-dominant trees in the stands, at lease 1O(
meters apart. Leaf and cone specimens were collected from five distinct as-

DONAHUE AND Lopez Upton, A new variety of Pinus greggii 1085

! . Summary of the sampling of Pens a sites included in the leaf, cone and seed morphol-
ogy, pe production and terpene chemistry stuc

Latitude-Longitude | Elevation Number of trees sampled
(m) I c
Southern populations
Cerro Perico, Hidalgo 20°44' N — 99°02' W 1830-1970 14 i) 24
Laguna Atezca, Hidalgo 20°49' N — 98°46' W 1250-1420 15 1S 16
Laguna Seca, Hidalgo 21°02' N —99°10' W 1670-1830 12 12 0
E] Madrono, Queretaro 21°16' N— 99°10! W -1650-1730 15 15 17
San are Queretaro 20°56' N — 99°34" Wo 2310-2380 15 15 15
Valle Verde, Queretaro 21°29'N —99°12' We 1150-1250 15 15 0
total 86 87 72
Northern populations
Cerro Perico, Hidalgo 20°44' N—99°02' W 1830-1970 14 1S 24
Cafion Los Lirios, Coahuila 25°22' N — 100°29' W 2260-2460 14 15 0)
Mesa del Rosario, Coahuila 253°26' N — 100°28' W 1920-2325 2 12 13
Santa Anita, Coahuila 25°27' N — 100°34' W 2515-2620 18 18 20
La Tapona, Nuevo Leon 24°43' N— 100°10' W 2090-2350 l i) 24
Las Placetas, N o Le 24°55'N — 100°11 2370-2520 13 15 22
Loma El Oregano, Coahuila 25°22' N — 100°55' W 2310-2350 14 15 19
total 86 90 98

a = leaf characters b = cone & seed characters ¢ = terpene analysis

pects in the crown, and leaves were sampled only from first-order branches,
prior to elongation. The five cones collected per tree were used in both the
morphology and seed production studies. Stem-xylem oleoresin was col-
lected from 170 trees for analysis of terpene chemistry using standard pro-
cedures. Details of the sampling procedure and chemical analysis for the
terpene study are given in Donahue et al. (1995).

Leaf and Cone Morphology
For the morphology study ten leaf characters were analyzed: number of needles
per fascicle, fascicle sheath length, needle length and width, number of ventral
and dorsal surface stomatal lines, number of stomata per 3 mm length on
dorsal surface, number of medial and internal resin canals and total num-
ber of resin canals. Ten needles per tree were measured for length, width
and number per fascicle (1720 needles total). Five needles per tree were analyzed
for the resin canal and stomata information (860 needles total). Five fas-
cicle sheaths per tree were measured for length (860 sheaths total). Leaf width
and stomatal counts were taken at the mid-point of the leaf, as well as the
resin canal assessment, using standard microtechniques.

The ten cone characters measured were cone length and width, cone scale
apophysis height and width, seed length and width, seed wing length and
width, seed coat thickness and seed weight. Five cones per tree were mea-

1086 Stipa 18(4)

eon

sured for length and width. Cone length was measured in a straight line

from the base to the top; cone curvature was excluded. Cone width was the

average of two perpendicular measurements taken at the widest point of
the cone. Cone scale apophysis height and width were measured on four
sides, perpendicular to each other, located at the widest part of the cone.

—

Seed wing width was measured at the widest point of the wing.

The leaf and cone data were analyzed using statistical procedures of the
Statistical Analysis System (Donahue & Lopez Upton 1996) First a multi-
variate stepwise discriminant analysis was performed on the data to deter-
mine which traits would be most useful in separating trees from the two
regions. A canonical discriminant analysis was conducted on the variables
selected by the stepwise discriminant analysis to look at patterns of differ-
entiation among populations, and calculate spatial Mahalanobis distances.

Analyses of variation by region were done on each individual variable to
determine which varied significantly between the northern and southern
groups of populations. Next a multivariate analysis of variation was con-
ducted on the subset of variables that were significantly different between
the two regions. Means and coefficients of variation were calculated and a
correlation analysis was conducted on all morphological traits.

Seed Production

For the seed production study, five cones per tree were assessed (885 cones
total). The extracted seeds were counted and classified as filled, empty, first-
year aborted, second-year aborted, or insect-damaged. The number of fer-
tile cone scales was counted to calculate seed potential and determine seed
efficiency. Of the seed production criteria, the number of fertile cone scales
is least affected by environmental variation. Population and individual tree
means and coefficients of variation were computed for each seed-yield trait
(Lopez Upton & Donahue 1995). An analysis of variance was conducted on
the individual tree means with a nested model: trees, populations and populations
within region (north and south regions). Waller-Duncan comparisons were
performed on population means to detect differences among them, and to
look for trends across regions.

Terpene Chemistry

The terpene chemistry analysis was performed on one resin sample from
each of 170 trees by gas chromatography (Donahue et al. 1995). The chemical
components were identified by comparison with known chemical standards,
and also compared to results from gas chromatography/mass spectrometry
analyses run on one sample from each of nine populations in the study. The
chemical compositions were reported as “percent of terpenes.” This included
all the monoterpenes and longifolene, a sesquiterpene, which was of par-

—

ticular interest. Since myrcene and carene were not separated, a combined

DONAHUE AND Lopez Upton, A new variety of Pinus greggil 1087

amount was reported for both. Population means were calculated for six
terpenes. To characterize individual trees as “high” or “low” types for specific
terpenes, frequency distributions were examined to help establish thresh-
old values as criteria for determining the point of separation for the con-
centration classes. Threshold values were established at points where the
trees’ frequency distributions might be separated into two modes.

Herbarium specimens examined

Of the specimens collected for and examined in the comprehensive study,

the following vouchers, which are now considered variety australis, are lo-
cated in the indicated herbaria:

MEXICO. Hidalgo: Laguna Atezca, 26 May 1993, Donahue & Lopez Upton F85 (A) F84
(F) F77, F90 (MICH) F76, F80, F83, F86, F89 (MO) F75, F81, F82 (NCSC) F78, F8&7
(TEX). Laguna Seca, 24 May 1993, Donahue & Lopez Upton D56 (A) D50 (F) D47 (MICH)
D49, DS1, D57, D55, D53 (MO) D54 (NCSC) D46 (TEX). Queretaro: Mpio. Landa de
Matamoros, Ejido El Madrofio, 19 May 1993, Donahue & Lopez Upton B27 (A) B20 (F) B19
(MICH) B18, B28, B26, B22 (MO) B23 (NCSC) B17 (TEX); San Joaquin, 17 May 1993,
Donahue . ee Upton Al, Al3 (A) A2, All (F) A5, A9 (MICH) AG, A8&, A10, Al2, Ald
(MO) Al i (NCSC) A3, A7 (TEX); Valle Verde, 22 May 1993, ee & Lopez Upton C44
(A) C4. 5 (E) C38 (MICH) C335, C37, C39, C42, C45 (MO) C36 (TEX).

The following vouchers from the comprehensive a are considered
variety greggi, and are located in the indicated herbaria:

MEXICO. Coahuila: Loma El Oregano, 5 Jun 1993, Donahue & Lopez Upton 1158 (A)
1156 (F) 1150 (MICH) 1153, 1155, 1157 (MO) 1152, 1166 (NCSC) 1146 (TEX); Los Lirios,
4 Jun 1993, Donahue & Lopez Upton H 139 (A) H136 (F) H132, H144 (MICH) 1134, H138,
H140, H143, H145, H148, H151 (MO) H135 (NCSC) 4131, H142 (TEX); Mesa del Rosario,
3 Jun 1993, Donahue & Lopez Upton G129(A)G107, G1 14 (F)G104, G112 (MICH) G/02,
G105, G108, G111, G115 (MO) G106, G113 (NCSC) G101, G110 (TEX), Santa Anita, 3
Jun 1993, Donahue & Lopez Upton G129 (A) G123 (MICH) G120, G122, G124, G127, G130
(MO) G119, G126 (NCSC) G/21 (TEX). Nuevo Leon: 2 See af Tie 993, Donahue
& Lopez Upton K184 (A) K128 (F) = 79, KIS88 (MICH) K , K180, K183, K186, K190
(MO) K181 (NCSC) K/78, K187 (TEX); Las Placetas, aa oe a & Lopez Upton
J170 (A) J167 (B) J164 ices) ies ey (MO) J163, J175 (TEX)

In addition to the 860 specimens from the comprehensive study, the authors
have also examined the following additional herbarium material:
MEXICO. Coahuila: Mountain sides, San Antonio near Saltillo, 30 Aug 1848, J. Gregg
402 (MO). Hidalgo: 8 km al este de Cardonal, Mpio. Ixmiquilpan, 21 Oct 1979, D. Cibrian
T. 2687 (CHAP); Rio Malila, Mpio. Molango. 1 Sep 1983 J., Lopez Garcia 359 (CHAP);
Molango. 7 Jan 1978, J. Perez Fitz 1258 (CHAP). NUEVO LEON: Cerro del Potosi, 20
km al oeste de Galeana, 2 Nov 1973, Castillo y Villar 1257 (CHAP); Cerro del Potosi, Mpio.
= aleana, 4 Mar 1978, T. Eguiluz 1261 (CHAP). Queretaro: Minas Viejas, Mpio. Zimapan,

i Jan 1978, A, res A, 1259 (CHAP); Carr. Jalpan-Xilita Km. 230, 2 Mar 1978, T:
ee 1260 (CHAP); Zona arqueologica Las Ranas, Mpio. San Joaquin, 20 Feb 1987, J.
Rzedowski 42515 “7 HAP).

During the course of several years of field study, the authors visited a
number of sites and examined material that was not archived in herbaria.

1088 Stipa 18(4

—

Table 2 summarizes details of those additional sites where trees of Pinus
gregeit were observed and information gathered for this work.

RESULTS AND DISCUSSION

The results from the morphology study given in Table 3 showed that
based on population means, the southern populations had significantly longer
needles, more stomata per unit of leaf length, lighter seeds and five times
greater frequency of internal resin canals than northern populations (Donahue
& Lopez Upton 1996). Although the number of leaves per fascicle did not
differ significantly between northern and southern trees, four-needle fas-
cicles did occur more frequently in the north. The total number of resin
canals did not differ between north and south, however trees from southern
populations had five times more internal resin canals. The values shown for
the northern and southern “regions” in Table 3 are means of the six indi-
vidual population means that were part of that region. The values shown
for (range) indicate the range of those six population means.

While both the northern and southern populations had the same num-
ber of lines of stomata on ventral and dorsal surfaces, the number of sto-
mata per 3 mm length of leaf was greater in southern trees. On average,
southern trees had narrower needles and wider seed wings. Northern trees
had thicker seedcoats and heavier seeds. In the morphology study, northern
and southern population means for needle length were significantly differ-
ent, without overlapping values.

In the seed production study, no traits were significantly different be-
tween north and south, however the number of fertile cones scales showed
a clear trend. Cones from southern trees had 28% more fertile cone scales
than those from northern trees. The number of fertile cone scales is consid-
ered to be under more genetic control than the other criteria.

Table 4 gives a summary of the terpene chemistry results. B-phellandrene
was found to be the single most abundant terpene in the chemistry analysis
of Pinus greget (51%) (Donahue et al. 1995). Southern populations had significantly
higher proportions of @-pinene and myrcene and lower proportions of li-
monene and longifolene than northern populations. Northern populations
had almost no variation in the frequency of high and low concentration types
of trees; all trees within a population were either “high” or “low”. In this
respect, trees from southern populations had varying numbers of high and
low concentrations of a-pinene, myrcene and limonene.Results of this study
indicated that genes that control these terpenes appearto be fixed in the
northern populations. Longifolene is the sole terpene that distinguishes trees
from northern and southern populations. Southern trees had a near-zero content
of longifolene, while northern trees had an average content of 5%.

DONAHUE AND Lopez Upton, A new variety of Pinus greggii 1089
; y &lCss

TABLE 2. List of additional sites observed but not included in the comprehensive study.

State Locality Latitude Longitude Elevation

Northern populations

Coahuila Agua Fria 25° 26'N 100° 30' W 2400
Coahuila Cafion de Caballos 25° 15'N 100° 55' W 2410
Coahuila Cerro El Potosi 24° 53'N 100° 13'W 2430-2500
Coahuila Jame 25° 21'N 100° 35' W 2450
Coahuila Puerto Chapultepec 25° 15'N 100° 56' W 2410
Coahuila Puerto Los Conejos 25° 28'N 100° 34' W 2380-2700
Coahuila Puerto San Juan 25° 25'N 100° 33' W 2630-2680
Nuevo Leon La Chona 24° 17'N 99° 58'W 2300
Nuevo Leon Ojo de Agua 24° 54'N 100° 12' W 2200
Southern populations

Hidalgo Cieneguilla 20° 44'N 99° 02' W 1860
Hidalgo E] Pinon 20° SO'N 99° 12'W 1830
Hidalgo Elochoxitlan 20° 45'N 98° 47' W 1710-1860
Hidalgo Jalamelco 20° 47'N 98° 42' W 1800-1950
Hidalgo Minas San Francisco 20° 48'N 99° 20' W 1950—2100
Hidalgo Molango 20° 50'N 98° 44' W 1400
Hidalgo Pemuxtitla 20° 49'N 98° 46' W 1400
Hidalgo Xochicoatlan 20° 50'N 98° 43'W 1840
Puebla Patoltecoya 20° 13'N 98° 03' W 1440
Veracruz Carrizal Chico 20° 26'N 98° 20' W 1580

2

Taste 3. Means by region and (ranges of means within region) and p-values for the characteristics
which varied significantly by geographic location.

Population Needle Needle #of #of Internal Wing Seed Seed Coat
(region) engt Width Stomata esin Width Weight | Thickness
(cm) (mm) Canals (mm) (mm)
Northern 11.3 i)

= 28 35.6 OL 5.95 1.73 0.146
(10.4-11.8) (1.19-1.35) (44.1—36.8) (0.0—0.18) (5.55—6.37) (1.61—1.87) (0.138—0.156)
Southern 12.5 1.18 38.8 0.48 6.4 1.47 0.138
(12.0-13.3) (1.09-1.24) (36.2—41.3)(0.28—0.91)(6.06-6.65) (1.18—1.58) (0.128—0.145)

Significance 0.002 0.016 0.01 0.002 0.012 0.005 0.037
level

weight of 100 seeds (gms)

Tas_e 4. A comparison of terpene composition (%) of southern and northern populations of Pinus

Leg.

Pinus greggit a-pinene B-pinene Myrcene/ Limonene (-phellandrene Longifolene
Carene

Southern populations 17 0.8 15 10 58 0.3

Northern populations 5 0.7 3 41 54 5)

1090 Stipa 18(4)

In summary, trees from northern populations of P. greggiz can be distin-
guished from southern ones using needle length, number of stomata, num-
ber of internal resin canals, seed wing width, seed weight, seed coat thick-
ness, number of fertile cone scales and terpene composition. Of these, needle
length, number of stomata, number of internal resin canals and percent content

on

of longifolene had non-overlapping population means in the studies.
Leaf Characteristics
Subsequent to the comprehensive study, the authors and colleagues made
informal observations of leaf color in native stands in Mexico, and crials
planted outside the species natural distribution both within and outside of
Mexico. Differences in leaf color were reported in exotic trials (Dvorak et
al. 1996), where color was noted as dark green for northern populations
and pale green for southern. Using Munsell color charts for plant tissues,
which assign a code based on the hue, value and chroma of an object, an
attempt was made to characterize the differences seen on live trees in native
stands in Coahuila and Queretaro, Mexico, and planted trials in Veracruz,
Mexico, and Louisiana, USA. Leaf color was found to be variable both within
and among trees, but variation could be generalized as differences between
hue classes. Northern populations were predominately hue class “S GY”,
value & chroma combinations 4/6, 4/8 and 5/6, while southern populations
ele prec ee hue class “7.5 GY”, value & chroma combinations 4/
, 4/6 and 5/6 (light green vs yellowish green). In addition to color difter-
ences it was observed that needles of northern trees are stiffer, more erect
than those of southern trees. Figure 2 is a photograph of two branches that
illustrate the differences in leaf color, length and rigidness between var. grege?/
and var. australis. The eighteen-inch long branches were collected from the
lower third portion of the crown of three-year-old trees planted in Singer,
Louisiana, USA. The trees were planted in field plots located adjacent to
each other, and are typical of the rest of the trees (approximately 400) planted
in the same plots. They are also representative of the differences in foliage

—

characteristics seen in the twelve native populations cited in this study.

Distinguishing characters for the two varieties are summarized in Table
5. Most of the quantitative characters reflect the statistical results presented
in Table 3, except for leaf length. Table 3 shows mean leaf lengths, whereas
Table 5 gives a range of values that takes into consideration individual leaves
and trees from our study.

Icther characteristics distinguish southern from northern populations of
P. gregert. In genetic field trials planted in Brazil, Colombia and South Africa,
progeny from southern populations grew significantly taller chan northern
progeny, and maintained their differences in foliage color and stiffness. Southern
trees planted in Brazil and Colombia were 100% taller than northern trees at

DONAHUE AND Lopez Upton, A new variety of Pinus greggii 1091

Tabe 5. Summary of the cl ; hat distinguish northern and southern populations of Pénws grege?i.
CHARACTER var. gregelt var. australis
Leaves
position erect frequently drooping
rigidness sciff flexible
color light green yellowish green
length 7-12 10-15
stomata number! 34-36 36-41
internal resin canals lacking sometimes | or 2
eeds
wing width (mm) 5.5-6.4 6.0—6.7
Terpenes (%)
a-pinene low high
limonene high low
longifolene high low
myrcene low high

‘ber 3 mm leaf length

three years of age, and in South Africa they were 17 to 30 % taller than northern
trees (Dvorak et al. 1996). Similar results were observed in field trials in Mexico,
where southern populations grew 106% taller at two years of age (Alba Landa
et al. 1998) and 118% taller at nearly three years of age (Lopez Ayala et al.
1999). Trees from southern populations showed greater resistance to pine pitch
canker disease caused by Fusarium subelutinans tsp. pini than northern trees
ina greenhouse screening trial (Hodge & Dvorak 2000). Although it has not
been demonstrated that the cause is genetic, flowering and pollination of P.
ereggii in Mexico occurs in April—May in southern populations, and in May—
June in northern populations (Lopez Upton & Donahue 1995).

Recent taxonomic works on Mexican pines by Perry (1991) and Farjon and
Styles (1997) did not recognize a distinction between the two populations
either because they did not observe a large enough sample size to detect the
differences, or attributed them to climatic effects. The differences in internal
leaf characteristics and terpene composition cited here are least likely caused
by environmental changes, while differences in leaf size and color remained
when the two populations were planted side by side in the same environment.

—_—

CONCLUSION

Based on the results of the aforementioned studies and observations, the
authors believe that trees from southern populations located in the states of
Hidalgo, Puebla, Queretaro, San Luis Potosi and Veracruz constitute a separate
taxon treated as follows:

1092 SIDA 18(4)
Pinus gress Engelm. ex Parl. var. australis Donahue & Lopez, var. nov.
ie igs. ne Type: MEXICO. Queretaro: Mpto. Landa de Matamoros, El Madrofio,
° 16' N, 99° 10°W, 1690 m, 19 May 1993, Donahue & Lopez Upton B30 (HOLO-

rypE: MO).

Varietas australis ab Pinus grevgt var. erect distinguibilis est foliis elongatis, flavovirentibus,

cum stomatibus plus per 3 mm longitudione, interdum Can ilibus resiniferis internis uno
vel duobus, et terpene compositis myrcene altis et longifolene demissis, et endemicis ad
Mexico centralis.

The variety australis has leaves in fascicles of 3, LO—-15 cm long, 1.0-1.3
mm wide, with 36—41 stomata per 3—mm leaf length. Resin canals are 3—A(--
6) in number, predominantly medial, and occasionally 1|—2 internal. Cone length
is 8-13(-14) cm and cone widths are 3—S cm. Seeds are S—7 mm in length
and 3—4 mm wide. Seed wing length is 11-16 mm, and width is 6-8 mm.

Pararypes: MEXICO. Queretaro: Mpio. Landa de Matamoros, El Madrofo, 21° 16'N,
99° LOW, 1690 m, 19 May 1993, Donahue & Lopez Upton B27 (A) B20 (FP) B19 (MICH)
B18 (MO) B23 (NCSC) B17 (TEX).

The variety greggu has leaves in fascicles of 3, 7-12 cm long, 1.2-1.4
mm wide, with 34—37 stomata per 3 mm leaf length. Resin canals are (2—
a in number, medial. Cone length is ohas 2 cm and cone widths are

—S cm. Seeds are 5-8 mm in length and 3-4 mm wide. Seed wing length
is 13-16 mm, and width is 5—7 mm.

ACKNOWLEDGMENTS

The research conducted in conjunction with this work was supported by
the CAMC ? Cooperative, North Carolina State University, USA, and
Colegio de Postgraduados en Ciencias Agricolas, Mexico. Thanks to James
Hardin (NCSC), Bill Dvorak (North Carolina State University, USA) and
Carlos Ramirez Herrera (Colegio de Postgraduados en Ciencias Agricolas,
Mexico) for their assistance with the research. Paul Fantz (North Carolina
State University, USA) gave generous assistance with the manuscript re-
view and Latin diagnosis. Special recognition is given to Jesse Parker Perry,
Jr., co whom the taxon is dedicated, for his assistance during many years of
fieldwork and research. Perry’s great enthusiasm and many years of accom-
plishments in the area has encouraged and influenced many others to re-
search and conserve the pine species of Mexico and Central America.

REFERENCES
ALBA LANDA, J., L. MENDIZABAL HERNANDEZ, and A. APARICIO RENTERIA. 1998. le
de un hoe de procedencia/progenie de Pinus gregeri Englem. en Coatepec, Veracrt
Mexico. Foresta Veracruzana 1:25—
ok 7 K. and J. Lopez Upton. 1996. Coase variation in leaf, cone and seed morphology
of Pinus greggi Engelm. in native forests. Forest Ecol. Managem. 82:145—

DONAHUE AND Lopez Upton, A new variety of Pinus greggii 1093

Fic. 2. A comparison of leaf characteristics of (A) Pinus greggii var. gregeii from La Tapona,
Nuevo Leon, and (B) Pinus greggii var. australis from El Madrofio, Queretaro. The trees are
growing in Louisiana, USA, produced from seed from the corresponding native stands in

Mexico.

DonanuE, J.K., J.P. Perry, A.E. SQuILLACE, and S. Liu. 1995. Geographic variation in stem-
xylem terpene chemistry in native populations of Pinus greggii Engelm. Forest Genet.
2:217-225.

Donanus, J.K. 1993. Geographic variation in Pinus greggit seedlings in relation to soil acidity.
In: Resolving tropical forest resource concerns through tree improvement, gene conser-
vation and comestication of new species. Proceedings, IUFRO Conference; 1993 O

t; Cali, aus Pp. 172-177.

ee W.S., E. KierzKa, and J.K. DonNaHue. 1996. Three-y
of Pinus greggit in the tropics and subtropics. Forest Ecol. Managem. ‘83: 123-

se on, A. and B.T. Strytes. 1997. Pinus (Pinaceae). Flora Neotropica. ee New

ork aeGec Garden, New

ee G.R. and W.S. Dvorak. 5600 . Pitch canker resistance of Central American and

exican pine species and Pinus radiata from Chile and New Zealand. New Forests (In

] a een } fy

Ss).

Lopez Ayata, J.L. 1998. Variacién intraespecifica en el crecimiento en alcura en el brote
terminal de eee nee Tesis ene Division de Ciencias Forestales, Universidad
Autonoma de Chapingo. Chay Oo

Lope Zz Upton, a and J K. Donanus, 1995, Sea production of P7us greggii in natural stands

ee Pl. Notes 46(3):8
Perry, 1P i a The pines of ee hae Central America. Timber Press, Inc. Portland.

Qa

1094 SIDA 18(4)
BOOKS RECEIVED

D. LayTen Davis and Mark T. NIELSEN (Editors). 1999. Tobacco: Produc-

tion, Chemistry and Technology. (ISBN 0-63 2-04791-7). Blackwell

Science Ltd., Osney Mead, Oxford OX2 OEL, UK. $164.95. 467 pp.
A collection of 30 agricultural and a eevee papers intended to document the status
of tobacco and its products today. The fourteen subject chapters include: 1) seed to smoke,
2) breeding and genetics, 3) eee uses and applications in tobacco improvement,
4) agronomy and physiology, 5) production practices, 6) major tobacco diseases, 7) tobacco
insect pests, 8) leaf chemistry, 9) physical properties of leaf tobacco, 10) marketing, pro-

cessing and storage, | 1) cigarette manufacturing, | 2) smoke chemistry, 13) cigars and cigarill

oOo =

14) smokeless eubaees:

P.J.M. Knippes. 1999. Growing Bulbs Indoors. ISBN 90-5410-4678).
A.A. Balkema, P.O. Box 1675, Rotterdam, NETHERLANDS (Fax:
31-104-413-4947), published in the United States by A. A. Balkema
Publishers, Old Post Rd., Brookfield, VT 05036 (Fax: 802-276-3837).
$25.00. 88 pp. text, 13 pp. glossary, appendices and index, 65 color
photos.

A horticultural guide to ornamental bulbs and their cultivation, including chapters on: 1)

main growing areas, 2) cultivation, 3) oul Ibs f from A to Z (alphé a ee - Be-

nus), 4) glossary, 5) bibliography, 6) cl “genera in terms ult

ty
7) classification of genera and species in terms of ornamental “ace. > useful addresses.

STANLEY D. JONES, JOSEPH K. Wiprr, and PauL M. MonrGomery. 1997. Vascular
Plants of Texas. A Comprehensive Checklist including Synonymy,
Bibliography, and Index. (ISBN 0-292-74044-1, hbk.). The Uni-
versity of Texas Press, P.O. Box 7819, Austin, TX 78713-7819, U.S.A
$55.00. 404 pp.

“This comprehensive list also inc ludes crops, persistent perennials, and naturalized plants.’

The table of contents includes: Preface, Acknowledgments, Introduction, Format, Scientific

Names: Species and Infraspecific Rank, Authors of Taxa, Common Names, Abbreviations
and Special Designations, Conserved Names, Cultivated Plants, Endangered and Threat-
ened Taxa, Federal Noxious Weeds, Hybrid Taxa, Misapplied Names, Orthography, State
Symbols, Synonyms, Unpublished Names, Summary of Taxa, Summary Table, Checklist,
Biblosiahy. and Index. A full review is forthcoming.

Stipa 18(4): 1094. 1999

THE GENUS MYRSINE (MYRSINACEAE)

IN VENEZUELA

JON M. RICKETSON

Missouri Botanical Garden

P.O. Box 299, St. Louis MO 63166-0299, U.S.A.
jon.ricketson@mobot.org; ricketson@ lehmann. mobot. org
JOHN J. PIPOLY III
Botanical Research Institute of Texas

509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.
ppipoly @ brit.org; clusia@latinmail.com

ABSTRACT

A synopsis of the genus Myrs7ve in Venezuela is provided. The generic description is
updated, along with discussions of its morphology and ecology in Venezuela. A key to the
species and subspecies is provided, along with descriptions, discussions of distribution,
ecology and conservation, and etymology for all species. Each of the 12 species known for
the country (one with two subspecies) is newly illustrated and its distribution is mapped.
Four binomials: Rae ambigna Mez in Engl (a oi ae of uid coriacea R. Br. ex

Roem. & Schult. subsp. coriacea), gat a Mez (M ina (Mez) Pipoly), Rapanea
nitida Mez 7 yr sine nitida (Mez) Pipoly), and Rapanea guianensis Aubl. (M. guianensis (Aubl.)
Kuntze) are lectotypified. Three additional binomials are relegated to synonymy.

RESUMEN

presenta una sinopsis del género Myrsime para Venezuela. Se actualiza la descripci6n
genérica, ara aee por discusiones sobre su morfologia y ecologia dentro del pats. Se
provee una clave para separar las especies y subespecies, descripciones, ehensiouc: sobre

distribucion, ae fa, conservacion ay e seiaolont Se ilustran y se ofrecen
de cada una de las 12 especies (1 1a con dos ae Se tse cuatro

binomios: ee ambigua Mez in Bask (= iii sine coriacea R. Br. ex Roem. & Schult. subsp.
coriacea), Rapanea andina Mez (=Myrsine andina (Mez) Pipol ee nitida Mez ( x yrsine
nitida (Mez) Pipoly), y Rapanea guianensis Aubl. (=M. cae (Aubl.) Kuntze) y tres especies
adicionales se relegan a la sinonimia.

INTRODUCTION

The genus Myrsine L. contains about 300 species, of which nearly one
fourth remain undescribed. C. Chen and Pipoly (1996), Pipoly (1991, 1992a,
1992b, 1996), Pipoly and C. Chen (1995), and Ricketson & Pipoly (1997)
have provided summaries of evidence for broader circumscription of the genus,
especially to include Rapanea Aubl. This circumscription has most recently
been accepted by Oregui (1998a) in her treatment of Myrszne for the South-
ern Cone of South America, and tentatively by Anderberg and Stahl (1995)

Sipa 18(4): 1095-1144. 1999

096 Sipa 18(4)

in their preliminary analysis of phylogeny in che entire order Primulales.
While Anderberg and Stahl (1995) were relatively noncommital regarding
the circumscription of Myrszne, later work by Stahl (1996) in determining
the systematic position of species formerly placed in Heberdenia Banks ex
DC., revealed that the genus would be paraphyletic if taxa formerly assigned
to Rapanea were removed from it, so the best circumscription of the group
is in its broadest sense. The conclusion was based on a cladogram using three
species from morphological extremes within the genus. Oregui (1998b)
constructed another cladogram ina paper discussing anemophily in the genus,
in which Myrs7ne was most closely related to the Papuasian endemic genus
F ee Mez. However, it should be noted that Fittingia is a group whose
species have been described from very eee material, and that there
has not been any analysis to include all genera in the family, so these cla-

dograms should be considered very tentative.

Myrsine is pantropically distributed, occurring in diverse vegetation types,
from mangroves to subalpine scrub, but always in moist, wet or pluvial habitats.
The genus is defined by be al (axillary), fasiculate or umbellate inflorescences,
sessile or on short, perennating peduncles girdled by persistent floral bracts
thus forming “short shoots.” In preparation for our treatment of the genus
for Flora of the Venezuelan Guyana, it became necessary to assemble speci-
mens from the entire country with complete synonymies and bibliographic
references, and to lectotypify several species. Because of the somewhat ab-
breviated format of that flora, the present synopsis is intended to provide a
complete nomenclator and a preliminary treatment for Flora de Venezuela
for this often misunderstood and nomenclaturally complex genus.

MORPHOLOGY

Habit and Architecture

The majority of Myrs/ne species in Venezuela are shrubs to small trees to 10
m tall, but occasional individuals of M. dependens (Ruiz & Pav.) Spreng., M.
cortacea (Sw.) R. Br. ex Roem. & Schult. subsp. coriacea, and M. guianensis
(Aubl.) Kuntze, have been known to reach heights of 15 m, 30 m, and 15
m, respectively. All species of Myrs/ve are terrestrial, with positively geo-
tropic roots, although those in cloud forests occasionally occur on deep humus
formed by decaying organic matter over large boulders.

All Venezuelan species of Myrsine, with the notable exception of M. dependens,
exhibit Rauh’s Model of architecture (Hallé et al. 1978), characterized by a
polyaxial, monopodial, rhythmically growing, readily distinguishable trunk,
that develops tiers of branches morphogenetically identical to itself. All branches
are orthotropic and monopodial, with spiral phyllotaxy and lateral (axil-
lary) inflorescences that do not affect shoot development. As was found for

RICKETSON AND P1poty, Myrsine in Venezuela LOOT

Myrsine cubana A. DC. (called M. floridana A. DC. in the paper), all branches
are sylleptic (Wheat 1980) producing two prophylls followed by adult leaves.
Myrsine dependens exhibits Massart’s Model of architecture (Hallé et al.
1978), characterized by a polyaxial, monopodial, rythmically growing, readily
distinguishable trunk, that develops regular tiers of branches at levels es-
tablished by the growth of the trunk meristem. Branches are plagiotropic
y leaf symmetry (distichous in this case), but never by apposition. The
position of the inflorescence is not significant in the definition of the model.
Like branching in Rauh’s Model, the branching is sylleptic. This model is
typical of many Myristicaceae, and is also known in Ardisia crenata Sims
and A. polycephala Wall. in the Myrsinaceae. Most species of the genus Embelia
Burm. f. also exhibit Massart’s Model. Symmetry of the entire plant is striking,
and allows immediate recognition of Myrszne dependens among the other species
in its subparamo thicket and paramo margin habitat, even though most
branches will fall due to lack of light in the compact canopy of the thicket.
With long-lived branches (when light conditions permit), Massart’s Model
is most often confused with Roux’s Model, differing only by its rythmic
(and not continuous or diffuse) branching. Field work conducted in Ven-
ezuela, Colombia, and Ecuador has shown that the branch tiers consist of
pseudoverticels, most often comprised of three branch units. It is notable
that Massart’s Model 1s typical in extreme environments, where wind-shearing
of orthotropic trunk axes produce the characteristic “Krummbholz Effect”
in taxa such as Abves balsamea Mill. (Hallé et al. 1978). That resilience in-
herent in Massart’s Model may be significant in the species’ ability to with-
stand extreme conditions of high winds, extreme diurnal temperature fluctuations,
soil compaction from overgrazing, and lack of oxygen due to high altitude,
all typical of the paramo margin and subpdramo thicket habitats.

Branchlets

A branchlet is here defined as the distal 10 cm of any branch. Branchlets in
Venezuelan Myrszne are straight, terete, smooth, glabrous, rufous or ferrugineous
villous- or floccose-tomentose, or rufous glandular-papillose, lenticellate or
not. Most species have thin outer bark, but in Myrsine guianensis, an extremely
active cork cambium results in significant cork accumulation, linked to dryness
of the habitat.

Leaves

Species of Myrsine, like all Myrsinaceae, are exstipulate and have simple leaves.
The leaves are mostly spiral, except in Myrsine dependens, where they are distichous.
In Venezuela, all species have petiolate leaves. A study of ptyxis (Cullen
1978) showed that four Asian and Pacific species of Myrsine surveyed had
supervolute ptyxis, but a survey among the Venezuelan species has not as

1098 Stipa 18(4)
yet been conducted, nor has the vernation been studied thus far. The leaf
blade texture may be chartaceous, coriaceous, thickly coriaceous or carti-
laginous. The difference between thickly coriaceous and cartilaginous 1s that
the leaf is rubbery and will not snap on bending perpendicular to the plane
of the midrib in the former, while it will easily snap with application of
moderate bending perpendicular to the plane of the midrib in the latter
The shape is most often obovate, oblanceolate, elliptic or oblong, but may
rarely be ovate (Myrsive maguireana Pipoly), or linear-lanceolate, narrowly
elliptic or lorate (M. ressnosa (A. C. Sm.) Pipoly). The apex may be attenu-
ate, acuminate, acute, obtuse, rounded, broadly rounded, or emarginate,
with or without a short, often blunt, bulbous, mucron formed by an exten-
sion of the midrib, while the base may be acute, cuneate, obtuse, or broadly
rounded, usually decurrent on the petiole. The adaxial surface may be gla-
brous, smooth or scrobiculate, with or without a villous-tomentum of uniseriate
hairs or glandular-papillae along the length of the midrib above; the sec-
ondary venation may be conspicuous, inconspicuous or not visible, and on
the lower surface, the punctation may be black or pellucid punctate or black
punctate and punctate-lineate; the margin is usually entire, and may be flat.
merely inrolled, or revolute at least basally. The petioles may be flat on the
adaxial surface, canaliculate, marginate or canaliculate and marginate dis-
tally, near the leaf blade base.
Inflorescence, Flowers and Fruit
The inflorescence in Myrsne consists of a sessile to subsessile umbel, or a
fascicle. The inflorescence bract is very early caducous and ts rarely seen on
herbarium specimens. The “peduncle” consists of an accrescent, perennat-
ing rachis that mimics a shoot, gradually built up by persistent, girdling
Horal bracts to form a “short shoot.” The floral bracts may be early cadu-
cous or persistent, chartaceous or coriaceous, orbicular, ovate, widely ovate,
deltate, obovate, apically acute, rounded or obtuse, inconspicuously pellu-
cid punctate, brown punctate, black or red punctate and/or punctate-lineate,
the margin most often entire, but may be erose and usually with glandular
cilia, but sometimes glabrous. In Venezuela, all pedicels are terete, and vary
mostly by their length and relative thickness. In most species the pedicel is
accrescent in fruit, but not exceeding approximately 5% of its original length.
The flowers are 4—5(—G6)-merous, perfect and normally functionally uni-
sexual. When they are bisexual, the androecium is smaller than that of a
staminate flower and the gynoecium is smaller than that of the pistillate
flower of the same species. Flower texture is chartaceous for all species in
Venezuela, except in Myrs/ne picturata, whose corolla is membranaceous. The
calyx may be coty

—_—
—_—

liform, or rarely, campanulate or cupuliform, with val-
vate lobes nearly free to fused 1/3 their length. The lobes may be triangu-

RICKETSON AND PipoLy, Myrsine in Venezuela 1099

lar-ovate, widely ovate, or deltate, apically rounded, obtuse, acute, acumi-
nate, short-acuminate or long-attenuate, long-acuminate, flat or prominently
keeled abaxially, inconspicously pellucid punctate, brown punctate, or densely
and prominently black punctate, and punctate-lineate, with margins en-
tire or apically erose or roughly dentate, glabrous, villous-ciliate, long-glandular-
ciliate, or glandular-ciliolate. The corolla may be cotyliform, campanulate
or cupuliform, except for Myrszne minima, where the nearly free lobes erect
in anthesis make it appear tubular. The lobes are valvate and may be linear-
lanceolate, lanceolate, ovate, elliptic, or oblong, apically rounded, obtuse,
subacute, sharply acute to attenuate, or acute. The ornamentation ranges
from inconspicuously pellucid punctate and punctate-lineate, densely and
prominently black punctate and punctate-lineate, or medially brown punctate.
The margins are usually entire, and glandular-granulose along its length.
The stamens and staminodes are similar but the latter are reduced in size.
The stamens are monodelphous (united into a tube by their filaments), and
the tube developmentally fused to the corolla tube, the stamens thus ap-
pearing epipetalous, or rarely are visible (MI. maguireana), or readily appar-
ent (M. minima, M. resinosa). The anthers may be broadly ovate, elliptic, or
oblong; apically acute to obtuse, and apiculate or not; basally cordate, sub-
cordate, deeply cordate, or sagittate, and the connective punctate or epunctate
dorsally. The antherodes may be lanceololate, obcordate, widely ovate, or
ovate, apically acute or obtuse and apiculate or not, basally sagittate or deeply
cordate, and the connective epunctate or punctate dorsally. The pistil may
be obnapiform or ellipsoid, or rarely conical (M. andina) or globose (M. resinosa).
The style in the bcc species is obsolete. The stigma may be morchelliform,
prismatic with 2—3 lobes, conic and spirally lobed with 4 lobes, or rarely
umbraculiform i” ‘vesinosa) with 4 vertically spirally twisted lobes, or lacinate
nitida) and 4-lobed. The eared is most often conic and hollow.
he fruits are usually globose to s se, or rarely ellipsoid (M. dependens)
or ae (M. picturata), prominent! ? black punctate and punctate-lineate,
or obscurely pellucid punctate, not costate or with inconspicuous longitu-

—

dinal costae.
ECOLOGY AND BIOGEOGRAPHY

Myrsine species, throughout their range, are most often associated with

moist or wet montane life zones. In Venezuela, one may find Myrsine dependens
through the Andes, mostly in paramo along the margins, at the upper limit
of the supdramo thicket habitat, but rarely also at the subparamo-cloud forest
transition zone as long as the site is open and exposed to wind. Where the
cloud forest life zone is well-defined, one may expect Myrsine andina in the
cloud forest just below the subparamo thicket, where M. coriacea subsp. coriacea

1100 Stipa 18(4)

may also occur in gaps and more disturbed places, along with Myrsine pellu-
cida. While Myrsine dependens and M. andina are less tolerant of disturbance,
M. pellucida is more tolerant, and M. cortacea subsp. coriacea is essentially a
“weed,”

en

Along the Atlantic coastal range and into the Guayana Region, Myrszne
guianensis occurs mostly in primary and secondary riparian forests, but also
rarely in premontane forests and upland tepui savannas. Below this life zone
in the eastern portion of the Guayana Region (Pantepui Floristic Province
sensu Huber 1995), Myrsine nitida is found in the same premontane forests,
Clusia-Magnolia, or Mora riparian forests. At the upper limit of that same
zone, Myrsine coriacea subsp. reticulata is found in the riparian formations.
Immediately below these Guayana formations, but in the drier savannas,
M. restnosa occurs.

Among the endemic upland tepui taxa, two species are endemic to Cerro
de la Neblina and vicinity, including Myrs/ne maguireana and M. perpauciflora,
while two others also known from Neblina, M. minima and M. picturata, are
also found in the easternmost area of Pantepui, near the border of Guyana.
Myrsine macrocarpa is restricted to the area around Cerro Huachamacari, in
Estado Amazonas, an area known for high numbers of endemics (Pipoly 1992c).

NOTES ON KEYS AND TERMINOLOGY

The keys are artificial and designed to expedite identification of herbarium
specimens. An attempt has been made to emphasize vegetative characters
to increase the keys’ usefulness with sterile material. The numbers appear-
ing before the taxa refer to their respective position in the key; any correla-
tions with phylogenetic relationships are coincidental. Quantitative and
qualitative data presented in keys and descriptions for floral parts and bracts
were taken from organs rehydrated from herbarium specimens by boiling
in water. Measurements from these range from 10% to 15% greater than
those measurements taken directly from dried material. Data regarding stem
diameters, inflorescence rachises, pedicels, leaf and fruit shape were taken
from dried herbarium specimens.

Description of general morphological features follows Lindley (1848),
Pipoly (1987, 1991, 1992a, 1992b, 1996), Pipoly and C. Chen (1995), and
Ricketson & Pipoly (1997) for the inflorescence, rachis pedicels and floral
parts. Description of leaf morphology follows Hickey (1984), trichome description
follows Theobald et al. (1984), and basic cell and tissue terminology follow
Metcalfe (1984).

TAXONOMIC TREATMENT

Myrsine L., Sp. Pl. 1: 196. 1753, Gen. Pl. ed 5: 90. 1754; Roem. & Schult.,
Syst. Veg. 503. 1819; A. DC., Trans. Linn. Soc., London, Bot. 17: 104.

RICKETSON AND Pipoty, Myrsine in Venezuela 1101

1834; Ann. Sci. Nat. Bot. 9: 292. 1841; Ann. Sci. Nat. Bot. 16: 65—
97, 129-196, t. 1-3, 8-9, 1841; A. DC. in DC., Prodr. 8:92. 1844;
Mig. in Mart., Fl. Bras. 10:306. 1856; Hook. f. in Benth. & Hook.,
Gen. Pl. 2:642. 1876; Mez in Engl., Pflanzenr. IV. 236(Heft 9):338
1902; E. Walker, Philipp. J. Sci. 73:184. 1940, Bot. Mag. Tokyo 67:249.
1954, Bull. Ag. Home Ec. Univ. Ryuku 2:76, Quart. J. Taiwan Mus.
12:164. 1959; Stearn, Bull. Brit. Mus. (Nat. Hist.), Bot. 4:174. 1969;
Fosberg & Sachet, Smithsonian Contr. Bot. 21:3—11. 1975; Lundell,
Phytologia 48:137. 1981, Phytologia 56:418. 1984; Pipoly, Novon
1:204. 1991, Caldasia 17:1. 1992, Novon 2:392. 1992; Pipoly & C.
Chen, Novon 5:360. 1995; Y.B. Harvey & Pipoly, Fl. Pico das Almas
487. 1995; C. Chen & Pipoly, Fl. China 15:34-38. 1996; Pipoly, Sida
17:115-162. 1996; Ricketson & Pipoly, Sida 17:579-589. 1997. Type

(by monotypy): Myrsine africana L.

Rapanea Aubl., Hist. Pl. Guiane 1:121. t. 46. ; A. Juissieau. Gen. Pl. 288. 1789;
iq. in Mart., Fl. Bras. 10:306. 1856; Mez in — , symb. Antill. 2:427. fee Mez
in Engl. ite unzenr. IV. 236(Heft 9):342. 1902; Pitard in LeComte, Fl. Gen. Indoch.

3(6): ae 1930 ere Publ. Field Mus. Nat. Hist. Ser 18(2):898. 1938; ae Fl

Madagascar 161:138. 1953; Lundell, Fieldiana, Bot. 24 (8-1):190. 1966, Fl. Panama

8:286. 1971; Fournet, Fl. Ilus. Phan. Guadeloupe et Martinique 1046. 1978; Taton,

Fl. Afr. Cent. Myrs. 53. 1980; Kupicha, Fl. Zambesiaca 7:201. 1983; Halliday, Fl.

stp. E. Afr. Myrs. 2:8. 1984; Little et al. Arbol. Puerto Rico y Islas Virg. 2:872.
1988 38

Diheandis Dombey ex lam., Encycl. 1:245. 1783.

Samara Sw., Prodr. 1:120. 1788, pro parte, non L.

Manglilla A. Juss., Gen. Pl. 151. 1

Arthrophyllum Lour., Fl. Cochinch. 1:120. 1790.

Caballeria Ruiz & Pav., Fl. Peruv. Prodr. 1:1 1794.

Roemeria Thunb. (non bi Nov. Gen. PI. 9:130. 1798.

Scleroxylum Willd., Ges. Naturf. Freunde on Mag. Neuesten Entdeck. Gesammten

Naturk. 3:57. 1809.

Suttonia A. Rich., Ess. Fl. Nouv.-Zel. 349. pl. 38. 1832.

Merista Banks & Sol. ex A. Cunn. in A. DC. in DC., Prodr. 8:95. 1844.

Shrubs or small trees. Leaves alternate, exstipulate. Inflorescences lateral (axillary)
umbellate or fasciculate, sessile or on short, perennating, accrescent peduncles
girdled by persistent floral bracts (thus forming “short shoots”). F/owers 4—
5(—6)-merous, bisexual or unisexual (the plants then bisexual, monoecious,
dioecious, or polygamous); sepals nearly free or united to 1/2 their length
imbricate or valvate, usually ciliate, punctate, persistent; petals nearly free
or rarely united to 1/2 their length, usually ciliate, glandular-granulose at
least along margin and often throughout within, punctate; stamens and
staminodes similar, subequalling corolla length, the filaments free or con-
nate basally to form a tube, the tube with or without sterile appendages
alternating with the filaments, and all merely adnate to the corolla tube; or

1102 Stipa 18(4)

developmentally fused throughout, the anthers or antherodes thus appear-
ing epipetalous, the anthers and antherodes similar, ovate or reniform, el-
liptic or oblong, rarely sagittate, 2-celled, dehiscing by longitudinal slits,
or rarely by subterminal pores opening later into wide longitudinal slits;
pistil and pistillode similar; conic, ellipsoid, obturbinate, obnapiform, or
variously subglobose; ovary globose, costate or not, glabrous or glabrescent;
ovules few, uniseriate, completely immersed in placenta or seated below apical
pores in placenta or variously projecting; style obsolete to present, taper-
ing into stigma; stigma morchelliform (morel-shaped), liguliform, sinuate
to lobate, prismatic and 3 (—4)-lobed, or rarely conical. Fraita globose, subglobose,
ellipsoid, ovoid, or obovoid or subovoid drupe, with somewhat fleshy exo-
carp and crusty or leathery endocarp, |-seeded; seed occupying cavity, the
endosperm horny, ruminate; embryo cylindric, transverse.

Myrsine, as here defined, contains ca. 300 species and is pantropically
distributed. In Venezuela, 12 species are known, separable by the follow-
ing key.

KEY TO MYRSINE OF VENEZELA

. Branchlets, petioles and/or leaf midrib densely ferrugineous or rufous vil-
lous-tomentose or Hoccose-tomentose, at times early glabrescent, the trichomes
uniseriate
2, iB anelies plagiotropic; leaves distichous, the blades (0.6—)1.5—2(—3.5) cm

ong, apically mucronate, scrobiculate abaya: the secondary veins not visible;

Howers 4(very rarely 5)-merous; ca

yx widely ovate to deltate, ets
without, the margins villous glandular-ciliate; — eee ieee
. Branches orthotropic; leaves apical, the blades (1. oN 3cm long, without

apical mucron, smooth above, the secot idary

—

M. dependens

i

raised at least adaxially; flowers miss 4)-merous; calyx tr ieacalersiyat

scattered papillose-| suberulent or glabrate without, the margins erose, glan-

dular-ciliate; fruit globos 2. M. coriacea.
3. Leaf blades 6—13 ¢m moe inflorescences (3—)5S—9(—1 1 )-flowered; caylx
lobes | a than wide 2a. M. coriacea subsp. coriacea
. Leaf blades (1.5—)3.5—5.5 (—5.8) cm long; inflorescences 2(—3)-flowered;
calyx lobes nee 2b. M. coriacea subsp. reticulata

Le Be eee petioles and leaf midrib glabrous or reddish glandular- papillose.
{ cee obsolete to 5 mm long.

. Leaf blades cartilaginous, (2.3—)3—5(5.4) cm wide; petioles th —3

mm diam.; calyx lobes ea! keeled, apically long-acuminate,

ee margins roughly det

&
nw

. M. maguireana
. Leaf blades coriaceous to othic kly coriaceous, 1—2(—2.9) cm wide; perils

thinner, obsolete to 1.5 mm diam.; calyx lobes flac, apically obtuse or

acute, the ses minutely erose or enti

6. Leaf blades (1.7—)2—3 cm long, 1-1.5 cm acre calyx lobes obtuse
apically, 7 margins minutely erose toward the apex, glabrous.

4. M. minima

RICKETSON AND Prpoty, Myrsine in Venezuela 1103

6. Leaf blades (2.8—)3—5.5(—6) cm long, 1.5—2.5(-2.9) cm wide; calyx

lobes acute apically, the margins entire, long glandular-ciliate to-

ward apex or densely glandular-ciliate throughout.
7. Branchlets red glandular ee tomentose apically; leaf blades

emarginate apically; pedicels 0.5—0.8 suoetans calyx lobe _
long glandular-ciliate ae Ps > apex . M. andina
7. Branchlets glabrous apically; leaf bl ae acute or rounded a
edicels 1.2—2 mm long; calyx lobe margins densely glandular-
ciliate shrougoe 5. M. perpauciflora
4. Petioles 5-20 mm long.
8. Leaf blades re. cm wide.
9. Leaf blades (3— a5 times s longer than wide, 4 —15 cm piIOne L=3(—

2.7)cm wide, lower |

lineate, the lines (S—)20— 40 1 mm long; peuoles flat hae: margin-

ate; pistillate pedicels 3.5—-5 mm long; calyx lobes apically

short-acuminate, the margins sparsely glandular-ciliolate; fruit

subglobose, longer than broad. 7.M. resinosa
9. Leaf blades 2—3 times longer than wide, (3—)4—6 cm long, 1.8-2(—

3) cm wide, lower leaf surface densely and prominently black poet

tate and very short punctate-lineate, the linesO.5—1 mm long; pet
oles canaliculate; pistillate pedicels 0.9-1 mm long; calyx lobes lly

long-attenuate, the margins glabrous; fruit obovoid 8. M. picturata
8. Leaf blades (2.7—)3—9.2 cm wide.
10. Pistillate pedicels obsolete to 1.4 mm long; secondary veins of the
leaf blades inconspicuous
11. Branchlets, ae ener leaf midrib glabrous or glandular-granu-
lose; pedicels 1.1—1.4 mm long; calyx lobes 1.2—1.4 mm long,
wider than long; fruits 8-12 mm in diam., obscurely pellucid
punctate. 9. M. macrocarpa
11. Branchlets, petioles and midrib of leaf blade reddish glandular-
papillose; pedicels obsolete to | mm long; calyx lobes 1 mm long,

longer than wide; fruits 3-3.5 mm in diam., obscurely on

tate and punctate-lineate. M. pellucida
10. Pedicels 1.5—3 mm long; secondary veins of the leaf Se con-
spicuous.
2. Leaf apices acute, rarely obtuse; lower leaf surfaces conspicu-
ously reddish tea and punctate-lineate; calyx lobes deltate;
fruit 3—3.5 mm diam., obscurely pellucid punctate; gallery, Mora
and Clusia- Magnolia f orests 11. M. nitida
12. Leaf apices obtuse to broad iy rounded; lower leaf surfaces in-

conspicuously black punctate to punctate- Linear, calyx lobes
y | |

ovate; fruit 3.5—5 mm diam. aii black ace; cloud
forests to lowland savann 12. M. guianensis

1. Myrsine dependens (Ruiz & Pav.) Spreng., (Figs. 1H, II, 1J, 3), Syst.

vee. ] oe 1825. Caballeria dependens Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil.

_ 1798. Manglilla dependens (Ruiz & Pav.) Roem. & Schult., Syst. Veg. 4:506.

: 9, is Ga (Ruiz & Pav.) Mez in Engl., Pflanzenr. [V. 236(Heft 9):377.

1902 4RU. HudAnuco: Near Mufia, without elev., H. Ruiz G J. Pavon s.n.
aaa core isoTyPes: F, G-DEL).

_

1104 Sipa 18(4)

\
a
Ps pes”
ec EE ES a

\

\

\
7%
AG

J

Fic. 1. A. Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult. subsp. coriacea, detail of branchlet
apex, petioles and midrib of leaf blades with dense tomentum, drawn from Pipo/y 6486. B
Habit, flo ce branch of same, drawn from FE. Breteler 4580. C. Fruit of same, doxea
from [ Breteler 4580. D. Habit, flowering ea of Myrsine coriacea (Sw.) R. Br. ex Roem.
& Schule, ue peace (Steyerm.) Pipoly, drawn from B. Magwire 33465. E. Fruit of
same, drawn from B. Magutre 33465. F. Abaxial leaf Sa same, drawn from B. Magwire 33465.
G. Aba leaf of same, oe eg drawn trom J. Steyermark 58983 (NY isotype).
H. Habit, flowering branch of Myrs/ oo v.) Spreng., drawn from L. Marcano-

Berti S04, I. Fruit of same, pave from af. Marcano-Berti 804, J. At aca leaf of same, drawn
om L, Marcano-Berti 804

RICKETSON AND Prpoty, Myrsine in Venezuela 1105

Myrsine ciliata Kunth in H.B.K., Nov. Gen. Sp. 3:248. 1819. Rapanea dependens (Ruiz &
Pav.) Mez in | var. ciliata (Kunth) Cuatrec., Rev. Acad. Colomb. Ci. Exact. 8:324.
1951. Type: COLOMBIA. Without locality, without elev., without date, A. von Humboldt

& A. Bonpland - oa (HOLOTYPE:

Samara myrtifolia Willd. ex Schule: & J.H. Schult. in Roem. & Schult., Mant. 3:220.
1 Myrsine myrtifolia (Schult. & J.H. Schult.) A. DC. in DC., Prodr. 8:103. 1844.
Tyre: COLOMBIA. Wichout locality, without elev., without ie A. von ears G
A. Bonpland s.n. (HOLOTYPE: P). According to TL-2, the third volume of Mantissa was
co- ae oe by Julius Herman Schultes, son of Josef August Schultes, in their revi-
sion of the Syst. Veg. that Josef and Johann Jakob Roemer had written earlier. There-
fore, we ~ changed the literature citation customarily used for the basionym ac-
cordingly.

Caballerta areola Rufz & Pav., ex A. DC. in DC., Prodr. 8:102. 1844. pro syn, nom.
inval. ERU. Without ae wih es without date, H. Ruz &_J. Pavon
s.”. (HOLOTYPE: G-DEL; Isorypres: G-BOIS A).
Rapanea pittiert Mez in Engl., Pflanzenr. IV. as 9):378. 1902. Myrsine pittieri (Mez)
Lundell, Phytologia 48:142. 1981. Type: COSTA RICA. Carraco: Volcan Irazti, 3,000
H. Pittier 14111 (HoLorype: B-destr., 1943; LEcroryPE: by Pipoly 1992b, BR;
ISOLECTOTYPE: US).
Rapanea peruv lana eee pecs acid: Ge LL ioe oe pernviana (Lundell) Lundell,
hytologia 48:1 1981. : PERU. Ct : Tres Cruces, upper edge of Parque
Nacional de ne 1-13 - NW of ee Pilcopata Road, 3,300—3,500 m,
29 Jun 1978, A. Gentry et al, 23478 (HoLoTyPE: LL-TEX; tsorypes: MO, USM).
Rapanea pittieri Mez var. chirripoensis Suessenguth, Bot. Jahrb. Syst. 72:281. 1942. Type:
OSTA RICA. SAN Jose: ae Grande, without elev., 28 Apr 1932, W. Kupper
1140 (HOLOTYPE: M; IsoTyPE: M).

Shrubs or small trees to15 m tall and 16 cm DBH; trunk orthotropic, the
branches plagiotropic. Branchlets terete, rufous puberulent or short villous
to densely villous-tomentose apically, at least when young, usually glabres-
cent, nodes congested, lenticels generally small and obscure. Leaves disti-
chous; blades coriaceous to cartilaginous, ovate to elliptic, obovate or sub-
orbicular, (0.6—)1.5—2.0(—3.5) cm long, 0.5—2 cm wide, apically rounded
or emarginate, with a short, often blunt, bulbous, mucron formed by an
extension of the midrib, basally obtuse to rounded, the midrib impressed
and puberulent to short-villous toward base above, prominently raised be-
low, the secondary venation not visible, nitid and scrobiculate above, bear-
ing numerous hydropotes, punctate and punctate-lineate below, the mar-
gin entire, densely punctate flat to inrolled; petioles canaliculate, 0.1—0.3(-0.4)
cm long, puberulent to short villous within, extending up the midrib of
the blade, outside often puberulent to short villous, usually glabrescent.
Staminate inflorescence fasciculate, 1—3(—4)-flowered; peduncles forming short
shoots 1—1.5 mm long, 1—1.2 mm diam., floral bracts chartaceous, oblate,
0.8—1 mm long, 1-1.5 mm wide, apically rounded, densely short-villous
above, glabrous below, the margins entire, densely glandular-ciliate; pedicels
1-1.8 mm long, glabrous. Staminate flowers 4(rarely 5 [Pipoly et al. 0466})-

1106 Sipa 18(4)

2 2?

merous, chartaceous, 3—3.5 mm long; calyx cotyliform, 1—1.2 mm, long,
the tube ca. 0.2 mm long, the lobes widely ovate to deltate, 0.8—1 mm long
and wide, apically acute, prominently black punctate and punctate-lineate,
the margin entire, villous-ciliate, especially toward apex; corolla campanu-
late, 2.5—3 mm long, the tube ca. 0.5 mm long, the lobes lanceolate to ovate,
2—2.5 mm long, 0.8—1 mm wide, apically subacute to obtuse, inconspicu-
ously punctate and punctate-lineate, the margins entire, minutely glandu-
lar-granulose; stamens 1.9—2.3 mm long, the filaments obsolete, the an-
thers broadly ovate to elliptic, 1.4—1.8 mm long, 0.4—0.6 mm wide, attached
dorsally at apex of corolla tube and thus appearing epipetalous, apically apiculate,

basally cordate, the connective epunctate dorsally; pistillode conic, hollow,
glabrous, the style obsolete, the stigma morchelliform, 0.1—0.2 mm long.
Pistillate inflorescence as in staminate but 1—3-flowered; peduncle 1—1.2 mm
long, 0.8—1 mm diam.; floral bracts 0.8—1 mm long and wide, densely cili-
ate along the margins; pedicels 1.7—2.2 mm, glabrous. Pisti/late flowers as
in staminate but corolla 2.4—2.6 mm long, the tube 0.2—0.4 mm long, the
lobes lanceolate, 2—2.2 mm long, 0.6—0.7 mm wide; staminodes resembling
stamens but abortive, the filaments obsolete, the antherodes lanceolate, 1—
1.4 mm long, 0.3—0.4 mm wide, apically apiculate, basally sagittate; pistil
obnapiform, the ovary subglobose to conical, glabrous the style obsolete.
the stigma morchelliform, 0.8—1 mm long, the placenta globose, the ovules

2

2-4, uniseriate. Frit ellipsoid, 3—3.5 mm long, 2.5—3 mm diam., densely

and prominently black punctate and punctate-lineate, with inconspicuous
longitudinal costae.

Distribution.—Myrsine dependens is known from Costa Rica and Panama,
and in the Andes from Venezuela to Bolivia, from 2,500—3,800 m eleva-
tion. In Venezuela, it is found in the Andean states of Mérida, Tachira, and
Trujillo, and along the coastal cordillera in and north of Caracas, in the Distrito
Federal and Miranda states. We expect it should also occur in the Serrania
de Turumiquire, in the states of Anzodtegui, Monagas and Sucre.

Ecology and conservation status.—Myrsine dependens is known from subparamo,
paramo margins, and subparamo-cloud forest transition areas, and with-
stands moderate to heavy disturbance. The principal obstacle to successful
seedling establishment appears to be compaction of the soil from overgraz-
ing by sheep and goats. The species is locally common, often forming dense
thickets, and is not threatened at this time.

Etymology.—The specific epithet comes from the latin “‘dependens’, meaning
suspended or hanging down, and refers to the often lax, narrow branchlets
that hang due to weight with age.

Specimens examined. VENEZUELA. Distrito Federal: Caracas, without elev., 1843
fl), H. Funck 481 (P); “crescit in declivitate meridionali montis Silla de Caracas, 800 hex,
without date (fr), A. vor Humboldt & A. Bonpland s.n. (P); Caracas, without elev., without

RICKETSON AND Pipoty, Myrsine in Venezuela 1107

date (fl), J. Linden 958 (K); Parque Nacional El Avila, Pico Naiguata, without elev., 5 Jan
1976 (fr), B. Manara s.n. (MO, NY, VEN); Cordillera del Avila, between Los Venados and
Pico Oriental, 1,675—2,640 m, 27—28 Feb 1944 (Al), J. Steyermark 55646 (F, NY); Silla de
Caracas, without elev., 1891-1892 (fr), J. Warmings s.n. (C). Mérida: Paramo de Mucuqut,
3,100 m, 7 Dec 1952 (fr), L. Bernardi 214 (G, NY, VEN); Near Alto del Aguada, 3,300—
3,500 m, 4 Nov 1976 (fr), L. Bernardi et al. 17136 (NY, VEN), 17144 (NY, VEN); Distrito
Libertador, Péramo El Escorial, vicinity of El Valle, NE of Mérida, ca. 2,800 m, 9 Sep 1982
(A), S. Clemants G J. Dugarte 2411 (F, NY, VEN); Distrito Miranda, road from es -
Aguila co Pifiango, at bridge, 3,400 m, 16 Sep 1982 (fr), S. Clemants G D. Diaz M. 2
(MERE, NY); Quebrada de en 3,220 m, 5 Apr 1930 (fr), H. Gehbriger 28 (F, MO, —
VEN); Paéramo near Hotel Los Frailes, N of the Mérida-Barinas Hwy, 08° 49! N, 70° 47!
O, 3,000—3,300 m, 29 Dec 1991 (fl), W Mezer 1097 (MO, VEN n.v.); Distrito Libertador,
Sierra de Culata, Ze kms NE of Mérida, 2,800—3,800 m, 19 Dec 1983 (fr), J. Pipoly et al.
6466 (MER, MO, NY); Dis trito Libertador, Sierra de Culata, 18 kms NE de Mérida, 2,800—
3,800 m, 19 Dec ae (fr), J. Prpoly et al. 6512 (MER, MO, NY); Distrito Libertador,
Parque Nacional Sim6n Bolivar, La Mucuy, Laguna del Coromoto, 3,400 m, 22 Dec 1983
(fr), J. Pipoly & G. Aymard 6566 (MER, MO, NY); Distrito Libertador, Municipio Tabay,
near Laguna de la Coromoto, Parque Nacional Sim6n Bolivar, 3,100—3,400 m, 19 Jun 1963
fl), L. Rugs ee 1669 (MERF, MO). vanes pecan of Fo de Naiguata, above Los
Chorros, 2,200—2,865 m, 16-17 Jun 1945 (fl), J. $ 7 (EF, MO). Tachira-Mérida
border: Parane. cn Negra, 3,000 m, 7 Oct 1965 (fr), J. Breer hie 9 (F, NY); Paramo La
Negra, ca. 3,080 m, 7 Oct 1965 (fl, fr), L. Marcano- a 804 (MER, MO). Trujillo: Paramo
Guirigay, couaid Pefia Blanca, 3,400 m, Aug 1958 (fl), L. Arig ie (NY, VEN);
Arriba de Jajf, Paramito, er Tufiame, rocky mountains, 3,1 Om, 29 Oct 1969
(fr), J. Cuatrecasas et al, 28185 (F); Along highway between F i. ee en and Bocon6, 64

m from Bocon6é, 500 m, Feb 1971 (A, fr), J. Steyermark 104785 (MO, NY). Without
location: Venezuela, without elev., 1842-1843 (fl), H. Funck 550 (BM); 1845 (A), H. Funck
& L. Schlim 58 (BM).

—

As here interpreted, Myrsine dependens includes a wide range of growth
forms, all restricted to Andean subpdramo and paramo habitats, extending
to the Caribbean coastal cordillera. The Myrsine ciliata growth form is the
most striking variant, chiefly because of its cartilaginous leaf blades with a
plicate habit. Further study is needed through the subparamos and jalca
habitat margins, from Colombia to Peru, to determine whether this vari-
ant deserves subspecific recognition

Among all Venezuelan species, Myrsine dependens is easily distinguished
because it exhibits architecture corresponding to Massart’s Model (Hallé et
al. 1978), characterized by an orthotropic trunk and plagiotropic branches
bearing distichous leaves. Fieldwork has shown it forms large populations
with high densities. The ellipsoid fruit is also distinctive. At this time, its
relationships are not certain, but the small-leaved taxa, including, M. mcrodonta
Pipoly, M. fosteri Pipoly, and M. brevis (J. F. Macbr.) Pipoly of Peru; M. lehmannii
(Standl.) Pipoly, and Myrsine paramensis (Cuatrec.) Pipoly of Colombia, seem
to form a monophyletic group defined by mucronulate to mucronate leaf
apices and Massart’s Model of architecture. All are high-altitude taxa growing
in harsh environments.

—

1108 Stipa 18(4)

2. Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult.

Shrihs or small trees to 8(—30) m tall; trunk and branches orthotropic. Branchlets
terete, ferrugineous to rufous villous tomentose to floccose tomentose, at
least apically, usually persistent, the trichomes uniseriate; lenticels small

and obscure to large and conspicuous. Leaves spiral; blades membranaceous,
chartaceous or conlareous, lanceolate, oblanceolate, obovate, elliptic or ob-
long, (1.5—)6-13 cm oe (0.8—)1—3 cm wide,apically acute or acuminate,
rarely obtuse to emarginate, basally acute, cuneate to obtuse or rarely rounded,

decurrent on the petiole, nitid and smooth above, pallid below, the midrib
flat or impressed above, prominently raised below, the secondary veins
prominulous to prominently raised above and/or below, conspicuously but
not prominently punctate and punctate-lineate below, villous at first, at least
along the midrib, glabrescent, the margins entire, flat or rarely revolute basally;

—

petioles canaliculate or marginate at least distally, 0.1—2 cm long, thick,
densely to sparsely ferrugineous or rufous villous tomentose, often glabres-
cent with age. Staminate inflorescence aa S—12-flowered; peduncles
forming short shoots |—4 mm long, 1—2 mm diam., glabrous or pilose; floral
bracts deltate to very widely ovate, 0.4—-0.8 mm long and wide, apically
obtuse to rounded, the margin glandular-ciliate; pedicels obsolete to 1.5
mm long, papillose-puberulent at first, glabrescent. Staminate flowers 5-merous,
very rarely with a few, scattered 4-merous ones on some inflorescences, 2.5—

A

4 mm long; calyx chartaceous, cotyliform, 0.4—1.5 mm long, the tube 0.1—
0.3 mm long, the lobes triangular-ovate, 0.4—1.2 mm long, 0.3—0.6 mm
wide, apically acute to obtuse, densely and prominently black punctate or
rarely epunctate, scattered papillose-puberulent without or glabrate, the
margins subentire to erose, glandular-ciliate; corolla chartaceous, campanulate,
2.5—4 mm long, the tube 0.5-1 mm long, the lobes lanceolate, 2.5—
mm long, 0.8—1.1 mm wide, apically subacute to obtuse, oe black
punctate-lineate, the margin entire, glandular-granulose; stamens 2—2.8 mm
long; filaments obsolete and anthers attached at the apex of earl tube,
thus appearing epipetalous; anthers oblong, |.5—1.8 mm long, 0.9—1 mm
wide, apically acute to obtuse, basally deeply cordate to subsagittate, the
connective epunctate dorsally; pistillode conic, hollow. Pisti/late inflorescence
as in staminate but oo mOweted, Pistillate flowers as in staminate but 1.8—
4 mm long; corolla 1.7—3.3(—3.9) mm; staminodes similar to stamens but
anthers ovate, 0.5—1 ee acuminate, basally slightly sagittate, the
connective epunctate; pistil obturbinate, the ovary 0.5—1 mm, globose to
subglobose; the style obsolete, the stigma morchelliform, (1—)1.2—1.4 mm
long; ovules 3, uniseriate. Frvit globose, 2.5—3.5(-4) mm diam., punctate to
punctate-lineate, glabrous, costa not prominent, brown or black at maturity.
As here circumscribed, Myrs/ne coriacea is the most widespread neotropical
species and perhaps the only “weedy” neotropical member of the Myrsinaceae.

RICKETSON AND Pipoty, Myrsine in Venezuela 1109

It is the most polymorphic of the ochlospecies (sensu White 1962; Prance
1982; Pipoly 1983) found in any myrsinaceous genus. While vegetative
plasticity is greater in this species than in any other in the family, the
morchelliform stigma is shared only with M. dependens. It is not certain whether
the morchelliform stigmas of both species are identical, or two states of what
could be a transformation series, and more study of this is needed. Among
the other distinguishing features of Myrsine coriacea are the few-flowered,
sessile, faciculate inflorescences, subsagittate anther and antherode bases,
the persistent ferrugineous tomentum of the vegetative organs and pedicels,
and the small, globose fruits.

2a. Myrsine ae (Sw.) R. Be. ex eae Se subsp. coriacea, (Figs.
A, 1 Ce 2). Syst. Veg. 4:511. 819. et coriacea Prodr. 1:32.
1788. a (Sw.) Mez in Urb. re Antill. 2:428. 1901. Type: JAMAICA
Without locality, without elev., without date, O, Swartz s.n. — BM).

Caballeria ferruginea Ruiz & Pav., Syst. Veg. Fl. Remy, ee 280. 1798. Manglilla ferruginea
(Rufz & og Roem. & Schult., Syst. Veg. 4:506. 1819. Myrsine ferruginea (Ruiz &
Pav.) Spreng., Syst. Veg. 1:664. 1825. Rinne fran (Rufz & Pav.) Mez in Urb.,
Symb. Antll. 2:429. 1901. Type: PERU. HuANuco: Near Mufia, without elev., without
date, H. Ruz et_J. Pavin s.n. (AOLOTYPE: i ISOTYPE: G).

Myrsine aha Kunth in H.B.K., Nov. Gen. Sp. 3:249. 1819. Type: COLOM-

ca: Near Popaydn, 1,000 m, without date, A. von Humboldt & A. Bonpland
1908 Ce P)

Myrsine jelskii Zahlbr., Ann. K. K. Naturhise. Hofmus. 7:3. 1892. Rapanea jelskii (Zahlbr.)
Mez in Engl. “Paanzene IV. 236(Heft 9):379. 1902. Type: PERU. Cajamarca: Near
Cutervo, Son elev. without date, C. von Jelski 15 (HoLotyPE: W; IsoTyPEs: KRA,

).

Samara saligna Willd. ex Schult. & J.H. Schult. in Roem. & Schult., Mant. 3:220.
Myrsine saligna (Schult. & J.H. Schult.) A. DC., Prodr. 8:103. 1844, Tyee: Herb, a
1039 (HoLoTyPE: B- WILLD, n.v.). According to TL-2, the third volume of Mantissa
was co-authored Julius Herman Schultes, son of Josef August Schultes, in their
revision of the Syst. Veg. that Josef and Johann Jakob Roemer had written earlier.
Therefore, we have changed the literature citation customarily used for the basionym

accordingly.

Myrsine eee -oides Schitdl., Linnaea 1833:525. 1833. Rapanea myricoides (Schitdl.) Lundell,
Wrightia 3:109. 1964. Type: MEXICO. Veracryz: Jalapa: without ae ity, without
elev., aa ut date, H. Galeotti 521, 522, 526 (syNtypes BR, FE ) fe

lectotypification ob this binomial until all material cited in the Sine and their
duplicates can be assembled

Myrsine tomentosa Presl, Reliq. Figen: 2:63. 1835. Type: PERU. HuANuco: without lo-

cality, without elev. without date, 7. Haenke s.n. (HOLOTYPE: PR).

Myrsine viridis Rusby, Mem. Torrey Bot. Club 6:74. 1896. Type: BOLIVIA. La Paz: Mapiri,
wichout elev., Jul-Aug 1892, M. Bang 1476 (HOLOTYPE:

Rapanea naling Mez in Engl., Pflanzenr. IV. 236(Heft 9):380. 1902. SYN. NOV. TYPE.

ELA. Disrriro Reon Caracas bei a Kolonie Tovar, without elev., with-

out en G. Karsten s.n. (Hotoryre: B-destroyed, 1943 (F Neg. # 4886); LEcro-
TYPE: here designated W (F Neg. # 31985).

Myrsine guatemalensis Gand., Bull. Soc. Bot. France 65:57. 1918. Type: GUATEMALA.

Stipa 18(4)

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Fic. 2. Distribution of Myrsne coriacea subsp. coriacea (*) in Venezuela.

RICKETSON AND Pipoty, Myrsine in Venezuela Py

Aura Verapaz: Coban, 1350 m, Dec 1906, H. von Tiirckheim 1001 (HOLOTYPE: P; ISOTYPES:
F, G, LL-TEX, MICH, M
— Lundell, Wrighta 5 a 1976. eas (Lundell) Lundell, he
81. Type: COST A. SAN José: Cordillera de Talamanca, Pacific
. Cae Massif, 2, a 5) Be m, 6 Apr ree C. Davidse & R. Pohl 1635 i
LOTYPE: LL-TEX; tsorypr: MO).

Mya vestita Lundell, Wrightia 7:274. 1984. Rapanea vestita (Lundell) Lundell, Phytologia
58:490. 1985. Type: COSTA RICA. Puntarenas: Cordillera de Talamanca, slopes sue
Cerro Echandf and Cerro cake 2,600—2 )m, 24 Aug 1983, G. Davidse et al. 2401
(HOLOTYPE: LL-TEX; tsorypes: CR n.v., ves

Myrsine microcalyx Lundell, os 58:277. 1985. Rapanea microcalyx (Lundell) Lundell
Phytologia 58:490. 1985. Type: PANAMA: Cuirigut: Cerro Colorado, 24 mi on gravel
from bridge over Rfo San Félix, 1,430 m, 22 Nov 1979, T. Antonio 2619 (HoLo-

E: LL-TEX; tsorype: MO)

Shrubs or small trees to 5(—30) m tall, 15(—50) cm DBH. Branchlets rufous
or ferrugineous villous- or floccose- tomentose, persistent at least apically,
rarely glabrescent. Leaf blades membranaceous, chartaceous or subcoriaceous,
6-13 cm long, 1-3 cm wide, the secondary veins prominulous to promi-
nently raised above, often conspicuously but rarely prominently punctate
and punctate-lineate, sometimes villous at first, especially along the mid-
rib on both surfaces, usually glabrescent with age the margins flat to revo-
lute; petioles 0.1—2 cm, thick, punctate, usually puberulent or short vil-
lous to densely villous-tomentose with reddish trichomes, often glabrescent.
Fruit globose, 2.5—3.5(—4) mm diam.

Distribution.—Myrsine coriacea subsp. coriacea is known from Mexico and
the West Indies through Central America, through the Andes from Ven-
ezuela to Argentina, and 1n montane or premontane areas in the Atlantic
coastal forest of Brazil, from 700—3,000 m elevation. In Venezuela (Fig. 2),
the species is known from the Guayana Region (Bolivar), the eastern states
of Anzoategui, Monagas and Sucre, the Coastal Range north of Caracas, including
Aragua, Carabobo, Distrito Federal and Miranda, and the entire Andean
region from Falcén to the Colombian border.

Ecology.—Myrsine coriacea subsp. coriacea occurs in primary and secondary
elfin, cloud, and wet tane (including montane tepui savanna), and subparamo
thickets. While no statistical analyses have been carried out, fieldwork has
shown that within a range from slight to heavy disturbance (mostly from
sheep grazing), Myrsine coriacea subsp. coriacea seems to increase in popula-
tion density with disturbance.

The subspecies’ range overlaps with that of subsp. reticulata only in the
state of Bolivar, where subsp. cortacea occurs in the talus slope forests, while
subsp. reticu/ata occurs in riparian gallery forests on the tepui summits. Given
its ability to thrive in disturbed habitats, subsp. corzacea it is not subject to
threat at this time.

i

11412 Sipa 18(4)
Etymology —The subspecific epithet, ‘coriacea’ refers to the coriaceous nature
of the leaf blades, an especially common feature in Caribbean populations.

Dae aie examined. VENEZUELA. Amazonas: oe Rio Negro, Cerro de
La Neblina, Expedition Camp VII, 00° 50' N, 65° 58! W, 1,850 m, 2 Dec 1984 (1), W.
Y, US

Anderson 1 3461 (EN S). Anzoategut: Disitrito Liberté i summit of Montanas Negras,
along the Sucre and Anzoategui woo 20 aa a NE of ea NE of Buenos
ires, Serranfa de Turimiquire, 10° 04' 30" N, ¢ 'W/, 2,000—2,350 m, 28 Nov 1981

(fr), G. Davidise G A. Gonzdlez 19540 (KF, MO, = hae (fl, fr), frre (MO, NY, VEN);
Cerro Corona, near radio installation, 2,000—2,200 m, 7 Jan 1987 (A), W. Hahn & FE. Grifo
ee (MO, US); Distrito Freites, Serranta - Turumiquire, Cerro Peonfa, ca. 10° 06'N,
06' W, 2,200—2,400 m, 8 Dec 1983 (fr), J. Pipoly 6450 (NY, VEN); Along forested
op of he: es of tributaries of Rio - vert, becween Rio Leén ans “Carmelita”, NE
of Bergantin, 800—1,200 m, 5 Mar 1945 (fl), J. Steyermark 61355 (F); Slopes of Montafia de
ie Palomas tributary of Rio Neveri, berween “Carmelita” and “Nata te ’, NE of Bergantin,
900—-1,000 m, 9 Mar 1945 (fl), J. Steyermark 61445 (F); S-facing steep slope, knife-edge
ridge above tree zone, Cerro Peonia (Cerro Los Paj oe above Santa Cruz, headwaters of
Rio Manantiales, E of Bergantin, 1,800—2,000 m, 20 Mar 1945 (A, fr), J. Steyermark 61615
(F). Aragua: Distrito Girardor, near Alto de C see 1,400 m, 11 May 1977 (fr), V: sages
7339 (MY, NY); Carretera las Tejerfas-La Tiara, highest point ee Carretera, 14-15 km S of
jancdan to Carretera Las Tejertas-Los Teques, 1,150—1,200 m, 4 Jan 1978 (fl), G. ae
& M. Pucci GO50 (NY); Summit, Pico Guacamaya, Banus Nacional Henri Pittier, 1,850
m, 25 Jan 1990 (fl), A. Cardozo & H. et 1218 (MO); Summit, Cerro Chimborazo,
Parque Nacional Henri Pittier, 2,230 m, 23 Feb 1990 (fl), A. Cardozo et al. 1278 (MO); On
steep slope, N slope of Pico Guacamay a, Parque Nacional Henri Pittier, 1,800 m, 23 Mar
1990 (fl), A. Cardozo et al. 13 O); Parque Nacional Henri Pitcier, without elev., 26
May 1990 (tr), A. Cardozo & ie Meneses 1421 (MO); Near Colonia Tovar, without elev.,
1854-1855 (fr), A. Fendler 758 (MO, NY); Distrito Ricaurte, after arch toward Colonia
Tovar, without ees 22 Jul 1987 (fr), C. de Rojas y F. Rojas 3682 (MY n.v., NY). Aragua:
Lagunita, Colonia Tovar, without elev., without date (fl),_/. Moritz 1175 (BM); ee of
Tovar, 1,200 m, 31 Jan 1928 (fl), H. Péttier 12793 (NY); Sabanas de Agua Negra, 1,500—
1,600 m, 18 Dec 1936 (fl), H. Pittier 13793 (US); S-facing slopes of Cordillera del Avila,
just above Caracas, along trail cowards Los Venados, 1,520—-1,675 m, 27 Dec 1943 (fl), J.
Steyermark 55022 (F). Barinas: Distrito Pedraza, SW of Carrizal, “La r scaza’on the S bank
of the Rio Canaguad, Parque Nacional Sierra Nevada, 08° 39' N, 70° 46' W, 26 Jun 1988
(fr), L. Dorr et al. 5693 (NY, PORT). Bolivar: Distrito Piar, Macizo del Chimanta, wide
valley in the SE sector of Apacara-tepui, in contact zone between Roraima a pe
extensive metamorfic (diabasic) intrusions, in the N sector of the Macizo 05°
O7' W, 2,150 m, 7 Feb 1984 (fh), M. Colella & O. Huber 451 rae NY); Distrito a
Kuaipia Min. Paso Con, 05° 37' N, 61° 46! W, ca. 1,000 m, 14 Apr 1986 (fr), 8. E/coro
y N. Vera 225 (MYF, NY); Reo Mun. Urdaneta, ene del Caront, “Sabanita”
LL km E of San hes O5° ( " N, 61° OO! 30" W, 1,120 m, 30 Jan 1985 (fr), L.
Herndndez y N. Dezzeo 114 ee ); Distrito Ping ca. 6 ee NE of Kavanayén, 05° 38'
N, 61° 40' W, 1,300 m, 6 Mar 1983 (fr), Huber & C. Alarcén 7375 (MYE, NY); Distrito
Aaa montane savannas ¢ ae ‘Piedra - nat ca. 3 km SE of Santa Elena de Uairén,
14° 35'N, 61° 06' W, 1,100—1,200 m, ie eG ),O. Huber & C, Alarcén 7931 (MYF,
— Distrito Piar, Macizo ef meee wide valley . ated in SE section of Apacard-tepui,

—

in the contact zone between Roraima Sandstones and extensive metamorphic (diabasic) in
the N sector of the Macizo, 05° 19'N, 62° 07! W, 2,150 m, 6—9 Feb 1984 (f), O. Hiber et

RICKETSON AND Pipoty, Myrsine in Venezuela 1113

al. 8786 (MYF, NY 2-sheets); - kms E of Kavanayén, ca. 1,200 m, 26 Jul 1983 (fr), R.
ral & A, Gonzdlez 70455 (MO, VDB); Entre Estacién y Aeropuerto CVG de Parupa, 1,250
m, 14 Sep 1983 ke G. Mowe et a 9606 (VEN); Gran Sabana, Via Kavanayén, El Jardin,
between Campamento Parupa and ana oe ca. 55 km E of Fuerte Luepa, without elev.
23 Jun 1983 (fl), N. Ramtrez 796 (VEN); Gran Sabana, Rastrojo, behind Campamento Rio
ae at junction with Rio Parupa, ae Nacional Canaima, without elev., 23 Nov 1993
,N. Ramirez et al. 4666 (MO); Ptari- ~~ steep ee slopes at base of first line of
cae bluffs, on S-facing part, E of “Cave Rock”, 2,130 m, 4 Nov 1944 (fl), J. Steyermark
59826 (F, MO, NY, US); Rio Karuai as savanna ate base of Ptari-tepui and
Sororopan-tepui, 1,220 m, 28 Nov a (bud), J. Steyermark 60737 (F),; Ridge above La
Laja at base of Sororopan-tepui, 1,375—1,460 m, 30 Nov 1944 (bud), J. Steyermark 60808
(F). Carabobo: Above Hacienda Cura, sake Valencia and dene 1 ae m, 8 Jan 1939
(fH), A. Adston 6172 (BM). Distrito Federal: Las Flores, Sierra de El Avila, 1,600 m, 15
Dec 1938 (fA), A. Adston 5515 (BM); Ecological study site, ae de ae Staal elev.,
29 Jan 1969 (fl), Z. Baruch 95 (NY); Bajo Seco, Facultad de Agronoméa Station, 2,000 m,
26 Jan 1983 (fl), L. C. de ena 3 ooh (BM); sae Nacional El] Avila, trail from end of

road to La Silla de Caracas, c ° 35'N, 66° 50' W, 1,951-2,316 m, 29 Jan 1984 (fl), J.
Luteyn et al, 9386 (NY, VE a ae Q387:F - ee Avila, 10° 32'8" oe a 53' W, 2,060
m, 11 Jun see (fr), W. Mezer 14 (MO); Cerro El Avila, S slope, al ridge t tel Humboldt-

Papelén, 10° 32' 03" N, 66° 52! o>” W, 1,930 m, 10 Jan 1992 (fl), 7 1317 (MO);
Fila del on 10° 32' 07" N, 66° 53' W, roe 2,100 m, 7 Jan 1992 (fr), W Mezer 1414

(MO); Headwaters of Quebrada Chacaito, 10° 33' N, 66° 52'O 2099 m, 3 Feb 1992 (fr),
W. Mezer eed (MO); Colinas W of Rio Marana ae 1,200 m, 4 mar 1971 (f1),G. Moriflo
592 (NY, VEN); 6 km ENE of Colonia Tovar, 1,900 m, 21 1960 (fl) J. Steyermark

86192 a VEN); Fila de Agua Negra, without elev., Feb 1938 (fl), EF Tamayo 436 (US,
VEN); Carretera de El Junquito, 1,700 m, May 1950 (fr), VEN Herb. No. 3879 (MO); Caracas,
without elev., 1891-1892 (fl), J. Warmings 553 (C); along Carretera de Los Flores y Boca
del Tigre, 1,600 m, without date (fl), L. Williams & A. Alston 314A (BM); Agua Negra,
1,400 m, 18 Mar 1938 (fl), L. Williams 9938 (F, US); Along road to Las Flores a Caligan,
1,700 m, 16 Dec 1938 (A), L. Williams & A. oo 10940 (EB). Falc6n: Cerro Santa Ana ,
Peninsula Paraguana, near top, 800 m, 15 Dec 1964 (fl), F Breteler 4289 (NY); Cerro Santa
a Peninsula Paraguand, 800-840 m, Dec ce (fl), 7 Lasser y L. Aristeswieta 3418 (F,
N); Peninsula de Paraguand, Cerro Santa Ana, isolated mountain, Nae elev., 17 Feb
iste (fl), G. Sobel et al, 2021 (NY); aes de a Luis, La Chapa y Uria, 1,400 m, 19 Jul
1967 (fl), J. Steyermark 99182 (F, MO, VEN). Falc6n and Lara: oe area be-
tween Falcén and Lara, Cerro Cerrén, on pat 1,800-—2,000 m, 27 Jun 1979 (fr), R. Liesner
et al. 8202 (MO, VEN), (fl), 8232 (MO, VEN); Cerro Socopo, 1,400-1,560 m, 29 Jun
1979 (fr), R. Liesner et al. 8389 (MO, NY, VEN). Lara: Distrito Moran, oes ae Humocaro
Bajo via Buenos Aires, 09° 36' N, 70° 03 W, 1,600 m, 13 Nov 1985 (fl,fr), H. van der Werff
&R, Rivero 78 38 (MO. NY); (fl), 7859 (MO, NY); Dini Moran, Carretera de Humocaro
Bajo, via Las Palmitas, without elev., 16 Nov 1985 (fl), H. van der Werff & R. Rivero 8009
Spee NY). Mérida: 10 km NE of Mérida, near village of Tabay, native coffee plantation,
1,900 m, 28 Oct 1963 (A), FE Breteler 3218 (NY); 19 km W of Mérida, 1,700 m, 31 Oct
1963 (fl), F Breteler 3242 (NY); 5 km NW of Mérida along road to El Valle, 1,920 m, 27
Feb 1964 (fl), F Breteler 3635 (NY); 10 km NE of Mérida, near village ‘Tebay 1 900 m, 9
a 1965 (fr), F es 4580 (MO, NY); NW of Mérida, on mee near Barrio Los Chorros,
1,950 m, 16 Jul 1966 (fr), J. de Bruijn 988 (MO); Valle Grande from the stream to the
paramo, 3,100-3, a m, 8 Nov 1976 ae s Charpin & F. i 13136 (NY); Vicin-
ity of Mérida, Rio Chama, without elev., 14 Jul 1951 (fr), H. Curran 2114 (NY); Distrito

1114 Stipa 18(4)

Rangel, Cuenca del Quebrada de La Mitistis, Parque Nacional Sierra Nevada, ca. 08° 51—
52'N, 70° 39' W, 2,400—2,750 m, 19 Jun 1988 (fl, fr), L. Dorr ape Barnett 5602 (NY),
Tabay, 2,200—2,300 m, 2 Sep 1930 (fr), H. Gebriger 401 (F, MO, NY); ceca Nacional
Sierra Nevada, de La Mucuy, 13 km air miles NNE of Mérida, 0 . 38) N, 71° 2' W, 2,400—
2,600 m, 9 May 1991 (A), W. Mezer & O. Carrero 869 (MO); Galipan, iMeneees Fac., with-
out elev., without date (A), J. Moritz 153 (BM); Distrito Libertador, Sierra de Culata, 18
km al NE de Mérida, 2,000—2,500 m, 19 Dec 1983 (fr), J. Pipoly et al. 6486 (MO, NY,
VEN); Distrito C ape Elias, Municipio Zerpa, Bosque Experimental de San Eusebio, 2,100—
2,400 m, 23 Jan 1963 (fl), L. Ruiz aa | 297 (MERE MO); Distrito Campo Elias, Municipio
Zerpa, El Molinillo- San Luis, between El ee and the village of La Azulita, ca. 1,680 m,
8 Mar 1963 (A), L. Razz Terdn 1436 (MERE, MO); Distrito Sucre, eet ae Estanques,
along Quebrada de Quirora, unos 4 km E of Quirora, 700—1,000 m, 2 | Feb 1970 (A), L.
Ruiz-Terdn & M. Lopez-Figuetras 120 (MERE, MO, NY); Estanques- oe de los Colorados
road, Mérida, 1,750 m, 12 Mar 1980 (fr), G. Sobel & J. Strudwick 2151 (NY); NW- & NE-
facing slopes above “La Isla”, above Tabay, 2,285—2,745 m, 18 May 1944 (fl D, bs aie
96588 (F); Distrito io. valley nee La Hechicera Mérida, 2,000—3,000 m
L985 (fr), E. Tanner & V. Kapos 107 (MO), Ridge behind la cere celete rien station, Ca.
2,600 m, 24 Nov 1985 (fr), E. Tanner & V. Kapos 246 (MO); 251 (MO); Spur ridge behind
Estacion La Montafia del Teleférico de Mérida, Tanner sites, a 0—2,750 m, 13 Jul Aa
(ster.), E. Tanner & V. Kapos 402 (MO); La Trampa, Carretera San Juan-Azulita, 2,500 m,
oe 1987 (fl), H. van der Werff et al. oe VEN), Disicrito Sucre, along the road ae
Santa Cruz de Mora to Canagua, along road 11.5 km from turn off paved road to G
08° 20'N, 71° 36' W, 1,660 m, 15 Dec a heat A. Weitzman & N. Holbrook 183 (MC
Y). Miranda: Altos de Pipe, 10° 23.7! N, 67° 0.1' W 1,500 m, 10 May 1963 (fl),
Agostini 174 (F, NY, VEN); Open sites near de San Antonio de los Altos, without elev., Feb
1965 (fl), L. Aristegnieta 5498 (MO, VEN); Reserva Biolégica, Instituto Venezolano de
vestigaciones Cientificas (VIC), Interior Branch of Cordillera de la Costa NE de Ven-
ae 10° 00' N, 66° 00' W, 1,700 m, 7 Jan 1987 (fr), N. Ramirez 2084 (NY, VEN).
ape ae Summit of Cerro de la ice eva de Dofia Anita, S of & bordering valley of Caripe,
1,300 m, 7 Apr 1945 (fr), J. Steyermark 61927 (F). Sucre: Peninsula de Paria, trail from
Los Pocitos de Santa isabel to Cerro Humo, 25 km NW de Irapa, 10° 41' N, 62° 36' W,
900-1, n, 12 Jul 1972 (fr), K. Dumont et al. 7611 (NY); Cerro Turimiquire, 10° 07!
53° 5 W. - O00—2,150 m, 14-16 Mar 1993 (fl, fr), W. Mlerer & G. Bronner 3523 (MO);
Pe ae e Paria, trail co Los Pocitos de Santa Isabel co Cerro Humo, 25 km NW de
Irapa, 10° 41' N, 62° 36' W, 900-1,250 m, 12 Jul 1972 (fr), G. Morillo 2626 (F, VEN);
Valley ee base of Cane de Diablo (W extention of S peak of Cerro Turumuquire),
and Cerro de Neverf, along headwaters of Rio de Amana, 2,000 m, 11 May 1945 (fl), J.
Steyermark 62721 (FE, ); Peninsula de Paria, Cerro de Humo, NW de Irapa, between
Roma and Santa Isabel, ca. 12 kms N of Rio Grande Arriba, 1,273 m, 2 Mar 1966 (fr), J.
Steyermark 94902 (NY, VEN); Distrito Marino and Distrito Arismendi, Peninsula c
trail between crossing of Rio Tacarigua to summit of slopes E of Cerro Humo, descending
to Las Melena N of Rio Grande Arriba, 10° 41' N, 62° 36—37' W, 760—1,000 m, 24 Feb
1980 (fA), J. Steyermark et al. 121737 (MO, NY). Pachira: Distrito Junin, entre Villa Paéz
y Betania, near Colombian border, 2,000—2,400 m, 15 Nov 1975 (fr), G. Bunting 4915
(NY); to above the right bank of Rio Tachira on the Colombian border, ca. 2,300 m, 13
Nov 1976 : A. Charpin & F- a aad 296 (NY); 7 km W of Rubio, 07° 42'N, 320
25' W, 900—1,000 m 18 Mar 1981 (fl), R. Liesner G A. Cis zdélez 10713 (NY, VEN); Slopes
at base of one de Tama, 2,47 . 2,550 m, 19 May 1967 (fl), J. Steyermark 98431 (MO,
NY, VEN); Forest of the small paramo of the Universidad Nacional Experimental del Tachira,
San Crist6bal, without elev., 10 Jun 1988 (fl), L. Valverde G L. Peta 1021 (MO); Distrito

panty

Qu.

e Paria,

RICKETSON AND Prpoty, Myrsine in Venezuela 1115

Uribante, ca. 5 km outside Siberia along old road to Pregonero, ca. 1,300 m, 20 Nov 1985
oe H. van der Werff & EF. ee pha ieee NY, PORT). Trujillo: Distrito Bocond, Guaramacal,

| E de Boconé, 09° N, 70° LL' W, 1,900—2,300 m, 7 Feb 1987 (fl), G. panel
et tah 5190 (F, MER, MO); en Bocono, Parque Nacional Guaramacal, 09° 15' 07" N

° 13'34" W, 1,950 m, 2-4 Jun 1995 (fl bud), N. Cuello et al, 931 (MO, PORT); N slopes:
a 14'48"N, 70° 12' 15" W, 1-3 Jul 1995 (ster.), N. Cuello 1107 (MO, PORT); S slopes,
09° 13' 32" N, 70° 10' O1" W, a OO m, 13-15 Dec 1995 (ster.), N. Cuello et al. 1256
(MO, PORT): 09° 12'45"N, 70° 09' 51" W, 2,300 m, 03-05 Jan 1996 (ster.), N. Cuello
et al, 1335 (MO, PORT), 1348 (MO, PORT): Distrito Carache, above Mesa Arriba, be-
tween ae Jab6n and Paéramo de Turmial, SE of Laguna de Turmal, 12 km SE of Carache,
09° 3 3° O9' W, 2,550—2,600 m, 12 May 1988 (fl), L. Dorr & L. Barnett 5157 (NY,
are oe (NY, PORT); Distrito Boconé and Disitrito Trujillo ai Serer La
Honda, SW of Arbol Redondo on the Boconé-Flor de Patria road, 09° 25
1 ,800—2,200 m, 2 Nov 1990 (fl), L. Dorr & - Barnett 7598 (MO, VEN); M oe ocon
Parque Nacional Guaramacal, S slopes, 09° 13' N, 70° 07! W, 2 100 m, 20— 52 Jan 1996
(ster.), A. Licata et al. 650 (MO, PORT): as Bocon6, Péramo Guaramacal, 4.4—4.8
kms beyond jct. NE of Boconé, 09° 15' N, 70° 14' W, 1,860 m, 19 Jan 1984 (fl), J. Lateyn
& J. Pipoly 9280 (F, MER, MO, ate Distrito Boconé, a Paéramo Arbol Redondo,
ca. 40 km N of Boconé, 09° 24' N, 70° 18' W, 2,073 m, 20 Jan 1984 (fl), J. Luteyn GJ.
Pipoly 9328 (MER, MO, NY), 9330 (MER, MO, NY); Distrito Carache, ca
Carache on Hwy. 2, 09° 38'N, 70° 09' W, 1,890 m, 21 Jan 1984 (fl), J. pan a Pipoly
9335 (MO, NY, VEN). Yaracuy: Distrito Nirgua-Distrito San Felipe border, Cerro La Chapa,
of Nirgua by road, 10° 12' N, 68° 35' W, 1,200-1,300 m, 21 Oct 1982 (A), G.

Davidse et al, 20813 (MO, NY, VEN); Sierra de Aroa, Cerro Negro, forest 8 km SW of San
Felipe, 10° 17' N, 69° 01' W, 1,200-1,800 m, 1-2 Apr 1980 (fr), R. Liesner & A. Gonzdlez
9933 (MO, VEN); El Amparo hacia Candelaria, a 7 km al N de Salom, 1,220—1,250 m,

—19 Jun 1972 (fr), J. Steyermark 106283 (NY, VEN). Without Location: Venezuela,
without elev., 1845 (fl), H. Funck & L. Schlim 140 (BM); Venezuela, without elev., without
date (fl), J. Moritz 360 (BM); Venezuela, 1,400 m, without date (fl), P. Vogel 177 (F); Ven-
ezuela, 1,400 m, 17 Mar 1946 (fr), P. Vogel 309 (EF).

As noted above, this highly variable species exhibits great variation in
quantitative features of its vegetative parts, and also in such features as the
degree of curvature of the leaf base (obtuse, rounded or acute), leaf shape,
vestiture thickness and trichome lenght, number of flowers and internode
length. All of these factors are responsible for the various segregates that
have been recognized in the past, including the new one synonymized above.
We believe we have assembled material from throughout subsp. coriacea’s
range for the first time since Mez’s work in 1902 and only for that reason have
we been able to appreciate how broadly variation occurs within and among
populations. Given this broad plasticity, we have adopted a very broad spe-
cies concept. One of the synonyms, Myrsine microcalyx, is a bisexual growth form
with consequent quantitative floral variation, but with qualitative features of
the vegetative organs within the normal range of variation for the subspecies.
2b. Myrsine coriacea (Sw.) R. Br. ex Roem. & Schultes subsp. reticulata

os es (Figs. 1D, LE, LF, 1G, 6), Novon 1:210. 1991. Rapanea
rm. , Fieldiana, Bot. 28: 477.1 1953. Type. VENEZUELA. Botivar: Mount

L116 Sipa 18(4)

Roraima, SW-facing forested slopes between Rond6n Camp and base of sandstone
— 0—2,255 m, 30 Sep 1944 (fl), J. Steyermark 58983 (HOLOTYPE: F eg.
): soTyPrs: NY, US, VEN).

Z!
~~

Shrubs or small trees to 3 m tall. Branchlets densely rufous villous-tomen-
tose, often glabrescent. Leaf blades coriaceous, obovate to oblanceolate, (1.5—
3.5—5.5(—5.8) cm long, (0.8—)1—2.5 cm wide, apically emarginate or ob-
tuse, basally acute, decurrent on the petiole, nitid above, pallid below, essentially
glabrous or with a few scattered hairs over the midrib above the petiole,
the margin revolute toward the base; petioles marginate, 4-5(—8) mm long,
densely villous along the margin, early glabrescent. Fruit globose, 3—3.5
mm diam.

Distribution.—Myrsine coriacea subsp. reticulata (Fig. 6) is endemic to the
Guayana Region, Pantepui Floristic Province in the state of Bolivar, Ven-
ezuela, growing at 2,000—2,750 m elevation.

Ecology and conserv ticulata occurs in gallery forests
along streams on the summits of tepuis, where it forms considerable stands
with several species of Ternstroemia, Bonnetia, Clusia, Gleasonia and Cybianthus
quelchii. Given the protection afforded the tepuis in the state of Bolivar, the
only threat to it is occasional fire. Therefore, it is not considered threat-
ened.

Etymology.—The subspecific epithet, ‘reticulata’ refers to the prominent
secondary veins of the leaf.

Spe scimens examined. VENEZUELA. Bolivar: Distrito Piar, Macizo del a wide
valle in the SE section of aes yacara- ke 05° 19' N, 62° 07! W, 2,150 m, 7 Feb 1984 (fl),
M.C loll & O. Huber 405 (M Y); Distrito Cedeno, Sierra de Maigualida, ae sector,

tepul plateau over c

jon

3

Rio Erebato, ca. LOO km directly SW of the Campamento Entrerfos, 05° 33'N, 6
2,100 m, 18 Nov 1988 (fl), O. Huber & L. lzguterdo 12802 (MYF, US): Te tepul, ms ran
Sabana, 7,000—8,000 fe [2,134—2,438 m], 17 Mar 1952 (fr), B. Maguire 33465 (F, NY,
US); Gran Sabana, Sororopan- tepul, near the C.V.G. Antenna, Parque Nacional Canaima,
without elev., 17 Aug 1993 (fr), N. Sete oe 4452 (MO, VE ve omen E
branch of ee ae of Rro Tirica, 2,15 200 m, 12 Feb 1955 (fr), J. Steyermark oe
Us urdack ue BRIT, FE, NY); Chimanta wee Torono-tepur, NW- nee forest
it Camp and base of escarpment, | ,880—1,970 m, 27 Feb 1955 (A), J. eae
Ey Wurdack 1194 (F, MO, NY).

issected caine at the headwaters of the Rio Chajura, Ww — es the
TW,

—
—
eo)

g

Subspecies reticulata is poorly known, but easily distinguished from subsp.

coriacea by its shorter leaf blades, fewer-flowered inflorescences, deltate ca-

lyx lobes, and riparian gallery forest habitat.

a ~ rsine maguireana Pipoly, oe 3, 4A, 4B), Novon 1:204. 1991.
. VENEZUELA. Amazonas: Cerro de La Nebli ina, Rio Yatua, Bonnetia forest
cs - Canon Cae 200—2,200 m, o Dec 1957 (tr), B. Maguire, J. Wurdack &
C. Maguire 42318 (oLoryee: VEN; tsorypes: MO, NY 3-sheets, US).

RICKETSON AND PIPoLy, Myrsine in Venezuela

L117

°
N

eee Meee

°
° °
i

°
oO

COLOMBIA

a

Myrsine picturata
Myrsine dependens
a= Myrsine andina

v
e

O= Myrsine maguireana

2
N

T T T : T
YNVANS HLIM GALAdSIG VIYV
LS
= ro]
Lu 34
= \
=
a
m0
ie. 7
Qe —
cag N
O <6
og
‘ [aa]

Distance in kilmometers

Fic. 3. Distribution of Myrsine picturata (VY ), M. dependens (*), M. andina (\)and M. maguireana

(OQ) in Venezuela.

1118 Stipa 18(4)

Fic. 4. A. Habit, flowering branch of Myrsine maguireana Pipoly, drawn from B. Maguire et
al. ey (NY isotype). B. Fruit of same, drawn from B. Maguire et al. 42 ene
C. Habit, flowering branch of eke ae nen ) Pipoly, drawn from R, Liesner 23107

me iflov

D. Fruit of same, drawn from R. Lieswer 23107 abit, flowering branch of Mfj per}
Pipoly, drawn from M. Nee 30697 (US isotype). F. Fruit of same, drawn from M. ‘Nee 30697
(US isotype). G. Habic, flowering branch of Myrsine andina (Mez) I -ipoly, drawn from _/,
Steyermark et al, 100998. H. Fruit of same, drawn from J. Steyermark et al. 100998.

RICKETSON AND Pipoty, Myrsine in Venezuela 1119

Shrubs to 1.5(—2) m tall; trunk and branches orthotropic. Branchlets ter-
ete, 3-7 mm diam., glabrous. Leaves spiral; blades cartilaginous, elliptic to
ovate, (4—)5—9 cm long, (2.3—)3—5.4 cm wide, apically and basally obtuse,
decurrent on the petiole, nitid above, pallid and scrobiculate below, the midrib
impressed above, prominently raised below, the secondary venation obscure,
conspicuously black punctate and punctate-lineate, the margin membra-
nous, hyaline, revolute, prominently black punctate, glandular-ciliate at first
when young, glabrescent, entire; petiole canaliculate, 0.4—0.9 cm long, 2—
3 mm diam., glabrous. Staminate inflorescence: unknown. Pistillate inflorescence:
an umbelliform glomerule, 4—9-flowered, the peduncle glabrous, epunctate,
1.7—2.5 mm long, built up by a series of floral bract bases; floral bracts obovate,
(.5—0.6 mm long, 0.9—1 mm wide, apically obtuse, the margins entire, densely
glandular-ciliate; pedicels cylindrical, 1-1.5 mm long, accrescent in fruit
to 2 (—2.5) mm long, glabrous, brown punctate. Pisti/late flowers 5-merous;
calyx chartaceous, cupuliform, erect, translucent, unequally divided, 1.6—
1.8 mm long, the tube 0.6—0.8 mm long, the lobes ovate, 1.1—1.3 mm long,
0.4—0.5 mm wide, apically long-acuminate, glabrous, medially brown punctate,
and prominently keeled, the margins hyaline, roughly dentate, highly ir-
regular, glabrous; corolla chartaceous, campanulate, 2.8—3.2 mm long, trans-
lucent, the tube 0.8—1 mm long, the lobes linear-lanceolate, 2—2.4 mm long,
1.2-1.4 mm wide, asymmetric, apically acute, medially brown punctate,
the margin glandular-granulose; staminodes 1.9—2 mm long, the anthers
appearing epipetalous, filaments thin, hyaline, 0.1—0.3 mm long, the antherodes
widely ovate, 1.2—1.5 mm long, 0.9-1.0 mm wide, the apically apiculate,
basally deeply cordate, the connective epunctate; pistil obnapiform, ca. 1.5
mm long, 1 mm diam., the ovary 0.8 mm long, costate, densely pellucid
punctate, glabrous, the style 0.2 mm long, costate, the stigma prismatic,
0.5 mm long, 2—3-lobed, the placenta hemispherical, the ovules 3, com-
pletely imbedded. Frit globose, 3-5 mm long and diam., densely pellucid
punctate.

Distribution.—Myrsine maguireana (Fig. 3) is known from Cerro de la Neblina
and Sierra de Maigualida, Amazonas, Venezuela, growing at 1,200—2,200
m elevation.

Ecology and conservation status. —Myrsine maguireana is restricted to dense,
open marshy scrub plateaus and on open tepui summits. While it has a highly
restricted distribution and narrow ecological tolerance, the remoteness of
these areas has thus far ensured that it is not threatened.

Etymology.—The specific epithet honors the late Bassett Maguire, inde-
fatigable explorer, collector and student of the Guayana Region's flora.

Specimens examined. VENEZUELA. Amazonas: Departamento Rio Negro, Cerro de
La Neblina, camp 9, 1.2 km NE of waterfall on E headwaters of Rfo Mawarinuma, 35 km
E of Base Camp, 01° 00' N, 65° 53' W, 1,780-1,820 m, 2 Feb 1985 (fr), B. Boom et al.

bo
=~
©

Stipa 18(4)

5535 (NY, US, VEN); Cerro Neblina Valle de Titricd, N of Pico Phelps, 00° 56' N, 65°

58' W, 2,200 m, | Dec 1984 (fr), 7) Croat 59551 (MO, VEN); Departamento Atures, Si-

erra de Maigualida, NW sector, dissected granitic mesa on the Heady otels of Rio Iguana,

tributary of the Rio Ventuari, 05° 40' N, 65° 08' W, 2,150 m, 24 Nov 1989 (fr), 0. Huber
13071 (MYE, MO), Cerro de La Neblina, Rio Yatua, E escarpment of upper Cafion Grande,

summit, La 2 2005 , 13 Dec 1957 (tr), B. Maguire et al. 42393 (NY 2-sheets); 14 Dec
1957 (fr), B. 3. Maguire oe 42403 (F, MO, NY 2-sheets, US, VEN).

Myrsine maguireana appears to be most closely related to M. glazioviana
Warm., a taxon known from the Planalto of Brazil. However, Myrsine maguireana
is easily recognized by the glabrous leaves, glandular-ciliate leaf bud mar-
gins, longer, glabrous pedicels, costate ovary and glabrous corolla lobe margins.

4, Myrsine minima (Steyerm.) Pipoly, (Figs. 4C, 4D, 6, SH ae ie enh
LOOL. Rapanea minima Steyerm., Ons Bot. 28:477. 19 . VEN
ELA. BoLivar: Summit, Cerro Roraima, 2,620—2,740 m, ie ar é ee. oe
Sreyermark 58848 os F; isoryprs: NY, VEN).

—

Shrubs 1.8—2.8 m tall; trunk and branches ortotropic. Branchlets terete,
glabrous. Leaves spiral; blades thickly coriaceous, obovate, widely oblong
or suborbicular, (1.7—)2—3 cm long, 1—1.5 cm wide, apically obtuse to broadly
rounded, basally obtuse to broadly rounded, nitid above, pallid below, the
secondary veins not visible, bearing numerous hydropotes in pits and con-
spicuously black punctate below, the margins scarious, flat, decurrent on
the petiole; petiole marginate, 3-6 mm long, essentially glabrous. Stami-
nate inflorescence a fascicle — flowered; pedicels 0.9-1.2 mm long. Stami-
nate flowers chartaceous, 3—3.2 mm long; calyx cotyliform, chartaceous, 0.9—
1.1 mm long and ee the tube ca. 0.1 mm long, the lobes very widely
ovate to suborbicular, 0.9—1.1 mm long and wide, apically obtuse, promi-
nently black punctate, the margins entire except minutely erose apically,
hyaline, glabrous; corolla cotyliform, 3—3.2 mm long, the tube 0.2—0.3 mm
long, the lobes oblong, 2.8—3 mm long, 1.2—1.3 mm wide, apically acute
to obtuse, cucullate, prominently black punctate abaxially, che margins densely
glandular-granulose throughout; stamens 2—2.2 mm long, the filaments 0.5—
0.6 mm long, the anthers oblong, 1.3—1.5 mm long, 0.7—0.8 mm wide,
apically apiculate, basally cordate, deshiscent by extremely wide longitu-
dinal slits, connective epunctate dorsally; piscillode lageniform, hollow, not
differentiated into ovary and style, 1.3—1.5 mm long, 0.3—0.4 mm wide.
Pistillate inflorescence as in staminate but (2—)3—5-flowered, pedicels 0.7—1
mm long. Pésti/late flowers as in staminate but 2—2.2 mm long; calyx coryliform,
hyaline, 0.8—1.1 mm long, the tube ca. 0.1 mm long, the lobes very widely
ovate to suborbicular, 0.9—1 mm long and wide, apically obtuse, promi-
nently black punctate, the margin entire except minutely erose apically, hyaline,
glabrous; corolla appearing tubular, but lobes nearly free, 2.0—2.2 mm long,
the tube 0.1—0.2 mm long, the lobes oblong, 1.8—1.9 mm long, 0.5—0.6

RICKETSON AND Prpoty, Myrsine in Venezuela L121

mm wide, apically obtuse to broadly rounded, densely and prominently black
punctate and punctate-lineate abaxially, densely glandular-granulose along
entire margin; staminodes 1—1.2 mm long, the filaments 0.7—0.8 mm long,
the antherodes obcordate, 0.8—0.9 mm long, 0.5—0.6 mm long, sterile, apically
acute, basally deeply cordate, the connective prominently brown puctate
apically; pistil ellipsoid, 1.7—1.8 mm long, the ovary 1.2—1.3 mm long,
().6-0.7 mm wide, opaque beige in color when dried, the stigma conic and
spirally lobed, 0.5—0.6 mm long, with 4 slight lobes; placenta ellipsoid,

4—0.5 mm long, 0.2—0.3 mm diam., bearing 3 uniseriate ovules, fully
exposed on the side of the placenta. Fruit globose, (2.5—)3—4.5 mm long
and in diam.

Distribution.—Myrsine minima (Figs. 6, 9) is endemic to the Guayana Region,
and is known from eastern Bolivar and adjacent Guyana, and from the Bra-
zilian side of Cerro de la Neblina, growing from 1,900—2,800 m elevation.

Ecology and conservation status. —Myrsine minima is known only from ex-
tremely remote and well-protected areas. It occurs in scrub forest and ex-
posed areas in upland tepui savannas. Therefore the species is not under
threat at this time.

Etymology.—The epithet “minima” refers to the diminutive size of the
leaf blades.

Specimens examined. BRAZIL. Amazonas: Parque Nacional do Pico da Neblina, rocky

formation of Pico da Neblina, 2,600 m, 21 Aug 1985 (fl), C. Farney et al, 905 (MO); Serra
da Neblina, summit to Pico Phelps, 9,000 ft [2,743 m}, 2 Dec 1965 (fr), B. Maguire et al.
60450 (BRIT, NY, US). GUYANA. Mazaruni-Potaro: Roraima, summit, La Proa camp,
E of border, near Lake Gladys, 05° 15' 36" N, 60° 13' W, 2,800 m, 14 Apr 1988 (fr), R.
Liesner 23296 (MO, US). VENEZUELA. Bolivar: Distrito Piar, Macizo del Chimanta,
north mesa of eae tepul, located in the SW sector of the masssif, 05° 10' N, 62° 16'
W, 2,200 m, 31 Jan 2 Feb 1984 Ca O. Huber & N. Dezzeo 8590 (MYF, NY); Distrito
Piar Macizo del Chimantd, sector SE, central SE section of Churi-tepui, 05° 15'N, 61° 58'
W, 2,250 m, 28 Me 1984 (fl), O. eee (MYE, NY); Distrito Piar, Macizo del Chimanta,
sector SE, mesa slightly sloping toward the SSE, in the central-SE portion of Churi-tepui,
05° 15' N, 61° 58' W, 2,250 m, 6-8 Feb 1985 (fr), O. ee et al. 1O100 (MYF, NY);
Distrito Piar, Macizo del Chimanta, NW sector, Supe pose mesa on the summit of the
central part of Murey-(Eruoda-) tepui, 05° 22' N, 62° 05' W, 2,600 m, 15-17 Mar 1986
(fl), O. Huber 11593 (MO, MYE, NY 2-sheets); Distrito Piar, W summit of Angasima-(Adanta-
) tepui, 10 km SSE from the W limit of Amurf-tepui (Macizo del Chimanté) and ca. 40 km
of the Mission of Wonkén, 05° 03' N, 62° 07! W, 2,100 m, 9 Aug 1986 (A), O.
Huber 11698 (MYF, NY); Kukendn-tepui, summit, 05° 13' N, 60° 18' W, 2,550 m, 11
Apr 1988 (fr), R. Liesner 23107 (BRIT, MO, NY, US, VEN); (fl, fr), 23270 (MO, US, VEN);
Ilu-tepui, lower plateau, 05°, 25' 36" N, 60° 29' W, 2,500 m, 16 Apr 1988 (fr), R. Liesner
23426 (BRIT, MO, US, VEN); Distrito Piar, Macizo del Chimanta, sector SE, Spee ace
sloping toward the SSE, in the central SW section of Churi-tepui, 05° 15'N, 61° 5
za m, 6-8 Feb 1985 (fr), J. Pipoly et al. 7141 (MO, MYF, NY, US); Chimanta ese
ntral portion of summit of Apdcara-tepui, 2,450—2,500 m, 21—22 Jun 1953 (fr), J.
eee 75931 (F, NY); Chimanta Massif, along tributary valley of E branch of headwa-
ters of Rfo Tirica, 2,120 m, 13 Feb 1955 (fl), J. Steyermark & J. Wurdack 843 (BRIT, F,

1122 Stipa 18(4)

ve Meseta,
nautical miles or GO kms NW of the mission at the medical camp of Rfo Kanarakuni,

1,922—2,100 m, 22- 27 Mar ae edi elas shee (MO, NY, VEN); Distrito Sucre,
Meseta de Jaua, 04° 35'N, ¢ 5' W, 2,020 m, ‘eb 1981 (A), J. Steyermark et al. 124328
(BRIT, MO, NY, ee Diserie Piar, Macizo ss . himanta, mesa at the southern base of
the upper walls of Apacard-tepui, N sector of Macizo, exposed knolls near stream, 05° 20!

, 62° 12! 2,200 m, 30 Janel Feb 1983 (fl), J. Steyermark et al. 128268 (MO, NY,
VEN); Roraima, 2,500 m, Jul 1910 (fl), E. Ule 8721 (US).

NY); Meseta de Jaéua, Cerro Jaua, summit of the Central-Western portion of t

>

Myrsine minima appears to be closely related to Myrsine andina, but is easily
separated by its glabrous branchlets, shorter leaves and longer pedicels.

5. Myrsine andina (Mez) Pipoly, (Figs. 3, 4G, 4H), Caldasia 17:3. 1992.
Rapanea andina Mez in Engl., Pflanzenr. IV. 236(Hetft 9):378. 1902. Type. ECUA-
DOR. Without location, without elev., 1857-1859, Rk. Spruce 5588 (LecroryPe, here
designated: K (pist. fl, fr); soLecrorypes: (pist. fl) BM, NY, (stam. fl) P-2 sheets, W-
2 sheets, a ae n.v.) FG, OXF)

Mez’ description clearly indicates that the branchlets of Myrsine andina are ferrugine-
ous-pilose tomentose, although they are often early glabrous. Also, the floral descrip-
tion matches those of all sheets except C. von Jelskz 12 (W). Given that Mez saw all
the duplicates of the Spruce collection, that they are che most numerous, and include
staminate and pistillate individuals, we designate its duplicate at Kew (K) as the lec-
totype

Shrubs or trees to 5 m tall; trunk and branches orthotropic. Branchlets ter-

ete, rufous glandular papillose-tomentose at first, glabrescent. Leaves spi-

ral; blades coriaceous, elliptic to oblong, (2.8—)3—5.2(—5.5) cm long, 1.5—

2.5(—2.9) cm wide, apically emarginate, basally obtuse to subacute, nitid,

the midrib impressed above, prominently raised below, the secondary ve-
nation not visible, densely and prominently black punctate above and be-
low, sparsely red-papillate along midrib above at first, early glabrescent,
the margin densely black punctate below, inrolled, entire, decurrent on the
petiole; petioles marginate, S—7 mm long. Staminate inflorescence fasciculate,

3—)5—8-flowered, on short perrennating shoots girdled by floral bracts, 1—

1.5 mm long; floral bracts ovate, 1—1.3 mm long, 0.5—0.7 mm wide, apically

acute, the margins somewhat crose, long glandular-ciliate; pedicels 0.5 mm

long. Staminate flowers 2—2.2 mm long; calyx chartaceous, cotyliform, the

tube ca. 0.1 mm long, the lobes ovate, 0.8—1 mm long, 0.5—0.7 mm wide,
apically acute, minutely and prominently black punctate, the margin en-

tire, long glandular-ciliate apically, glabrescent; corolla campanulate, 2—

2.2 mm long, the tube 0.2 mm long, the lobes ovate, |.8—2 mm long, 0.6—

0.7 mm wide, apically acute, densely and prominently black punctate and

punctate-lineate without, glabrous within, the margin entire, densely glandular-

ciliate apically; stamens 1.8—2 mm long, the filaments not discernible,
developmentally fused to corolla tube, the anthers appearing epipetalous,
broadly ovate, 1.5—1.6 mm long, 0.6-0.8 mm wide, apically apiculate, basally

RICKETSON AND Pipoy, Myrsine in Venezuela 1123

cordate, the connective slightly dark brownish dorsally; pistillode conic,
hollow or absent. Pésti//ate inflorescence as in staminate but on shoots to 1.5
mm long; pedicels 0.7—0.8 mm long. Pésti//late flowers as in staminate but
2.8—3 mm long; calyx cupuliform, 1.3—1.5 mm long, the tube ca. 0.3 mm
long, the lobes ovate, 0.9—1 mm long, 0.7-0.8 mm wide; corolla campanulate,
2.6-2.9 mm long, the tube ca. 0.5 mm long, the lobes 2.1—2.3 cm long,
0.6—0.8 mm wide; staminodes like the stamens but 1—1.3 mm long; pistil
conical, 0.8—1.0 mm long, 0.7—0.8 mm diam., the stigma sessile, conical,
pristmatic, with 4 lobes; placenta globose; ovules 2—3, uniseriate. Fruit globose,
3.5—S mm diam, the stigma persistent, prominently punctate.

Distribution.—In Venezuela, Myrsine andina (Fig. 3) is only known from
secondary and dry forests in Estado Tachira, but is well known otherwise
from Venezuela, southward through Colombia, Ecuador, and Peru to Bo-
livia, growing at (400—)1,200—2,500 m elevation.

Ecology and conservation status.—Myrsine andina is restricted to primary
cloud forest margins just below the subparamo thicket transition through-
out its range. Because many montane roads through the Andes follow the
paramo-cloud forest contour, it should be considered threatened.

Etymology.—The specific epithet refers to its place of description and range,
throughout the Andes

Specimens examined. VENEZUELA. Tachira: 5 km E of San Antonio del Tachira, 07°
N, 72° 25' W, 1,400 m, 3 Jan 1989 (fl), W. Hahn & F. Grifo 4971 (MO, NY, US);
Parque Cazadero, Quebrada Cazadero, 16 km NW of San Cristébal, 400-650 m, 2 May
an (fr), R. aes & M. Guarielia es (MO, VEN); Paramito between Quebrada de
Palmar & Quebrada de at base of Paramo de Tama, 2 kms above Betania & 7
kms above Villa Paéz, 2,500 m, 14 Jun 1944 (fl), J. Steyermark a a NY); Along path
between “Las Copas” nd peak a de Tierra Negra, toward C 1 Fundacion

Bélgica, on the narrow range er divides the headwaters of Rio Quinimari, Rio Frio and
Uribante and Rio Talco (Oird), 20—25 km S of San Vicente de la Revancha, 35—40 km S of
Alquitrana, SW of Santa Ana, le 2,880 m, 16 Jan 1968 (fr), J. Speirs etal. 100998
(NY, VEN). Without Locality: Without locality, without elev., 1848 (fl), J. Linden 108
(P)

Myrsine andina appears to be closely related to M. minima, but is easiliy
separated by its glandular-papillate branchlets, emarginate leaf blades and
subsessile flowers.

6. Myrsine perpauciflora Pipoly, (Figs. 4E, 4F, 9), me 2:176. 1992.
Type. VENEZUELA. Amazonas: Cerro de La Neblina, Camp VH, 5 km NE of Pico
Phelps, vicinity of heliport on rock outcrop, 00° 50! 40" N a or =H W, 1,850
m, 1 Feb 1985 (fr), M. Nee 30697 (HoLoTYPE: VEN; isotypes: F, NY, US).

Trees to 4 m tall; trunk and branches orthotropic. Branchlets terete, 2.5—
3 mm diam., glabrous. Leaves spiral; blades chartaceous, oblanceolate to elliptic,

(4—)4.8—5.5(—6) cm long, (1.5—)1.8—2 cm wide, apically acute, basally cu-

1124 Stipa 18(4)

neate to rounded, decurrent on the petiole, asymmetric, somewhat nitid
above, pallid and black punctate-lineate below, the midrib impressed above,
prominently raised and brown punctate-lineate below, the secondary veins
prominulous above and below, the margin revolute, translucent but not hyaline,
entire, brown punctate, glandular-ciliate at first, early glabrescent; petiole
marginate, 0.3—0.5 mm long. Staminate inflorescence. unknown. Pistillate inflorescence:
unknown. Infractescence: glomerulate, apparently 1—3-flowered, the peduncle
glabrous, epunctate, (1.2—)1.6-2.4 mm long; floral bracts orbicular, 0.8—1
mm long, 0.8—1 mm wide, apically obtuse, densely pellucid punctate, the
margins densely erose-ciliate; pedicels cylindrical, 1.2—2 mm long, glabrous,
sparsely pellucid punctate-lineate. Prsti//ate flowers: unknown. Fruiting ca-
/yx chartaceous, cotyliform, unequally divided, 1.2—1.3 mm long, translu-
cent, densely and prominently brown punctate and punctate-lineate, with
darkened glandular areas at the calyx lobe/tube sinus, the tube 0.2—0.3 mm
long, the lobes very widely ovate to deltate, 1—-1.2 mm long, 0.8—1 mm aide:
apically acute, prominently brown punctate and punctate-lineate, flat, the
margin hyaline, entire, densely glandular-ciliate. Frit globose, 3-5 mm long
and in diam., when dried, densely pellucid punctate and punctate-lineate.

Distribution.—Myrsine perpauciflora (Fig. 9) is endemic to the Cerro de La
Neblina, 1,500—1,730 m elevation.

Ecology and conservation status. — Myrsine perpauciflora is restricted to low
shrubland on saturated soils. Because Cerro de la Neblina is extremely re-
mote and protected, the species is not threatened.

Etymology.—The specific epithet refers to the fact that the inflorescences
are infrequent and very few-flowered.

Specimens examined. VENEZUELA. Amazonas: po ee Rio Negro, Cerro de

La Neblina, Cam} > VI, 00° 52' N, 65° 58' W, 1,730-1,850 m, 10 Feb 1985 (fr), 8. Renner
a (US); € ene La Neblina, Camp 2, Neblina massif, 2.8 = NE of Pico Phelps, 00°
49' 40" N, 65° 59' W, 2,100 m, 15 Apr 1984 (fr), B. Stein & A. Gentry 1525 (MO, NY,
EN)

Myrsine perpauciflora is most closely related to M. lancifolia Mart., but is
separated by the smooth upper leaf blade surface, Bes prominent brown
punctations of the calyx, and inflorescences with 1—3 flowers. Within the
Guayana Region, it may be confused with Myrszne ae. but is easily
recognized by the abaxial leaf surface less conspicuously (not densely and
prominently) black punctate-lineate, |—3 (not 3—5)-flowered inflorescence,
calyx lobes brown (not black) punctate, and globose (not obovoid) fruit.

re pa resinosa (A.C. pe Pipoly, (Figs. 5A, 5B 6), Novon 1:210.
1991. a edd resinosa A.C. Sm., Bull. Torrey Bot. Club 67:297. 1940. Tyrer. VEN-
EZUELA. Botivar: Rio ene near Arabapu, 4,200 ft [1,280 m], 17 Jan 1939

(A), A. Pinkus ae oe NY (pisc. fl.); lsorypes: F, GH, NY (2-sheets, US).

RICKETSON AND Prpoty, Myrsine in Venezuela 1125

Fic. 5. A. Habit, flowering branch of Myrsine resinosa (A.C. Sm.) Pipoly, drawn from R,
Liesner 23534. B. Fruit of same, drawn from R. Liesner 23534. C. Habit, flowering branch
of Myrsine picturata Pipoly, drawn from B. Maguire et al. 42375 (NY isotype). D. Fruit of

same, drawn from B. Maguire et al. 42375 (NY isotype).

Shrubs 1\-1.5(—3) m tall; trunk and branches orthotropic. Branchlets terete,
glabrous. Leaves spiral; blades chartaceous to thinly coriaceous, linear-oblanceolate,
lorate or narrowly elliptic, (4—-)6—-12(-15) cm long, (1—)1.7—2(—2.7) mm
wide, apically acute to attenuate, basally acute to att te, midrib prominulous
above, prominently raised below, the secondary venation prominulous above

1126 Stipa 18(4)

and below, nitid above, pallid below, with conspicuous black punctate-lin-
eations (S—)20—40 mm long below, the margins entire, inrolled and revo-
lute at least basally; petioles flat above, marginate, 6—9 mm long. Staminate
inflorescence a sessile, 4—8-flowered umbel; floral bracts ovate, 0.8—1 mm long,
0.6-0.7 mm wide, apically acute, inconspicuously red punctate, the mar-
gins entire, densely glandular-ciliolate; pedicels 2.5—3.5 mm long. Stamz-
nate flowers S(—7)-merous, chartaceous; 2.5—3 mm long; calyx cotyliform,
1.2-1.4 mm long, the tube ca. 0.2 mm the lobes ovate, 1—1.2 mm long,
0.9-1 mm wide, apica

—

ly short-acuminate, essentially epunctate or with one
or two scattered glands, the margin irregular, sparsely glandular-ciliolate;
corolla campanulate, 2.6—2.7 mm long, the tube 0.4 mm long, the lobes
ovate, 2.2-2.3 mm long, 1.6—-1.7 mm wide, apically sharply acute to at-
tenuate, sparsely brown punctate and punctate-lineate, the margins densely
glandular-granulose; stamens 2—2.2 mm long, the filaments not visible, the
anthers oblong, 2—2.2 mm long, apically apiculate, basally subcordate, the
connective dark brownish punctate-lineate dorsally; pistillode conic, 0.9—
| mm long, hollow. Pisti/late inflorescence a sessile S—7-flowered umbel, as in
staminate but pedicels 3.5—5 mm long. Pisti//ate flowers as in staminate but
2—2.2 mm long; calyx lobes ovate, 1.3—1.4 mm long, 0.9-1 mm wide; co-
rolla 2—2.2 mm long, the tube, 0.2 mm long, the lobes 1.8—2 mm long,
0.7—0.9 mm wide; staminodes |.3—1.5 mm long, the filaments 0.3 mm
long, the antherodes 1.1—1.3 mm long, 0.5—0.6 mm wide, apically apicu-
late, basally sagittate; pisctil globose, 2—2.2 mm long, the ovary |—1.1 mm
long, |.2-1.4 mmm wide, the stigma umbraculiform, with 4 vertical spi-
rally twisted lobes, 1—1.2 mm long; placenta globose, 0.6—0.7 mm long,
0.5—0.6 mm wide; ovules 2—3, uniseriate, immersed. Fruit subglobose, longer
than broad, 3.5—5 mm long and broad, black at maturity, the exocarp thick,
juicy, densely and conspicuously black punctate-lineate when dried.

Distribution.—Myrsine resinosa (Fig. 6) is known only from the eastern
portion of Pancepui Floristic Province in the state of Bolfvar in Venezuela,
and the adjacent Mazaruni-Potaro region in Guyana, growing at 470—1,000
m elevation.

Ecology and conservation status.—This species is known from gallery for-
ests along rivers in lowland dry savannas. Because these forests house the
relatively scarce source of construction materials, M. resinosa should be considered
threatened.

Specimens examined. GUYANA. Mazaruni-Potaro: Upper Mazaruni River basin, Kukui
River between Mokay River and Suru-agu-puh River, 470 m, 11 Sep 1960 (fl), 8. Tillett &
C. Tillett 45380 (FDG, NY, US). VENEZUELA. Bolivar: Gran Sabana, ca. 10 km SW of
Karaurin-tepui at junccion of Rio Karaurin and Rio Asadon (Rio Sanpa), 05° 19' N, 61°
03' W, 900-1,000 m, 20 Apr 1988 (fr), R. Liesmer 23465 (BRIT, MO, NY, US, VEN); 21
Apr 1988, 23534 (BRIT, MO, NY, US, VEN); Gran Sabana, orillas del Rio Aponguao,

RICKETSON AND Pipoty, Myrsine in Venezuela 1127
without elev., 18 Jan 1973 (fl), G. Morillo et al. 2887 (VEN 2-sheets); Along Rio Karuai,
between base of Ptari-tepui and Sororopan-tepui, 1,220 m, 28 Nov 1944 (fr), J. Steyermark
60726 (F, NY, US).

Myrsine resinosa may be confused with M. picturata, but is easily separated by the linear-
oblanceolate, lorate or narrowly elliptic leaf blades with long, conspicuous punctate-linea-
tions, and the longer pedicels.

8. Myrsine picturata Pipoly, (Figs. 3, 5C, 5D, 6), Novon 1:204. 1991.
Cafio Grande, 1,200—2,200 m, 13 Dec 1957 (A, fr), B. Maguire, oe Wurdack & C.
Waurdack 42375 (HOLOTYPE: VEN; isotypes: F, MO, NY 2-sheets,

Trees to 8 m tall. Branchlets terete, 4-5 mm diam. ae trunk and

branches orthotropic. Leaves spiral; blades coriaceous, elliptic to oblong, (3)

4—6 cm long, (1.5—)1.8—2(—3) cm wide, apically obtuse to slightly emar-
ginate, basally cuneate, decurrent on the petiole, the midrib strongly im-
pressed above, raised and ribbed below, the secondary veins prominulous
below, nitid above, pallid below, prominently and densely black punctate
and short-lineate, translucent glandular-lepidote above in bud, glabrescent;
the margin revolute, entire, hyaline, with prominently raised black punc-
tations, densely red glandular-ciliate in bud, glabrescent; petiole canaliculate,
(0.7—)0.8—1 cm long, glabrous. Staminate inflorescence. unknown. Pistillate
inflorescence glomerulate, 3—5-flowered, the peduncle glabrous, epunctate,
|.2—2.1 mm long; floral bracts membranaceous, very widely ovate to deltate,
0.7-1.0 mm long, 0.8—1.2 mm wide, apically obtuse, densely and promi-
nently black punctate and punctate-lineate, the margin entire, glandular-
ciliate; pedicels cylindrical, 0.9—1.0 mm long, glabrous, densely and prominently
slack punctate-lineate. Pisti//ate flowers 5-merous, translucent green; calyx
chartaceous, cotyliform, 1.3—1.5 mm long, the tube 0.1—0.2 mm long, the
lobes ovate, 1.1—1.2 mm long, 0.6—0.8 mm wide, apically long-attenuate,
glabrous, densely and prominently black punctate-lineate, medially crassate,
the margin hyaline, flat, irregular, glabrous; corolla membranaceous, cam-
panulate, 1.8—2.0 mm long, the tube 0.5—0.6 mm long, the lobes ovate,
1.2-1.4 mm long, 0.6-0.7 mm wide, apically rounded to obtuse, densely
and prominently black punctate and punctate-lineate, hyaline, the margin
densely glandular-granulose, entire; staminodia 1.2—1.4 mm long, the fila-

ments obsolete to 0.1 mm long, the antherodes ovate, 0.9—1 mm long, 0.3—

0.4 mm wide, apically apiculate, basally oe epunctate, but the con-

nective darkened dorsally; pistil ellipsoid, 2.1-2.4 mm long, the ovary 1.5—1.8

mm long, 1.0—1.2 mm diam., densely and ee black punctate and

punctate-lineate, the stigma subsessile, conical, ca. 0.6 mm long, 2—3-lobed

longitudinally, pellucid punctate, apically cuspidate, ovules 3, immersed

ina globose placenta. Frit obovoid, 4-5 mm long, 3—3.5 mm diam. when

dried, densely and prominently black pune and punctate-lineate.
Distribution.—Myrsine picturata (Figs. 3, 6) is endemic to Cerro de la Neblina

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Fic. 6. Distribution of Myrsine resinosa (mg), M. minima (*), M. coriacea subsp. reticulata (V),
M. nitida (K) and M. picturata (O) in Venezuela and bordering Guyana region. Note that

this map is essentially a close-up of the eastern portion of Figure 9, with M. minima re-
peated to eastern range more precisely.

RICKETSON AND Pipoty, Myrsine in Venezuela Li2)

of Amazonas and the [-tepui on the Gran Sabana of Bolivar, growing at
1,200—2,450 m elevation.

Ecology and conservation status.—This species grows in low woodlands in
cloud forests on saturated soil. Despite its resericted distribution, the pro-
tected status and remoteness of the Cerro provide it adequate protection.

Etymology.—The specific epithet refers to the dense and prominent black
punctate lineations of the leaf blades, perianth parts and fruits.

Specimens examined. VENEZUELA. a Departmento Rio Negro, Cerro de
La Neblina, Cumbre Camp Swale, 1,20( 00 m, 19 Nov 1957 (fr), B. Maguire et al.
42090 (FE, NY-2 sheets, US); Cerro de a a ee Yatua, S rim of upper basin of Cafion
Grande, 1,200—2,200 m, 13 Dec 1957 (fr), B. Maguire et al. 42376 (NY 3-sheets). Boliv
Gran Sabana, IlG-tepui, slopes below upper most W-facing escarpment, 7,000—8,0 00. fe
{2,134-2,438 m]}, 21 Mar 1952 (fl), B. Maguire 33535 (NY)

Myrsine picturata may be confused with M. perpauciflora, but is separated

y the more densely and prominently black punctate-lineate abaxial leaf
—5 (not 1—3)-flowered inflorescences, calyx lobes black (not

5

surface, the 3
brown) punctate, and obovoid (not globose) fruit.

9. Myrsine macrocarpa Pipoly, (Figs. 7A, 7B, 9), Novon 1:207. 1991.

Type. VENEZ UELA. AMAzZoONaS: Serrania Parti, Rio Parti, Cerro Part, valley above

ae and valley draining eastward through cumbre, 2,000 m, 10 Feb 1951 (fr), R,
Cowan & J. Wardack 313572 (HoLotyee: VEN; tsorypes: F, NY-2 sheets, US).

Tree to 7 m tall; trunk and branches orthotropic. Branchlets terete, 7-10
mm diam., glabrous. Leaves spiral; blades coriaceous, elliptic to obovate,
9.5-16 cm long, 6.5—9.2 cm wide, apically obtuse, basally acute, decur-
rent on the petiole, pallid and scrobiculate above and below, midrib im-
pressed above, prominently raised below, the secondary vein pairs 13-26,
mostly inconspicuous, but somewhat visible below, not evidently punctate,
the margin opaque, highly revolute basally, glabrescent, entire; petioles marginate,
1.5—2cm long, gla brous. Staminate inflorescence: unknown. Prstz//ate inflovesece Ce:
unknown. Infructescence: glomerulate, apparently 4—8-flowered, the peduncle
glabrous, epunctate, 1.2—4.2 mm long, made up of floral bract bases; floral
bracts very widely ovate to oblate, 0.5—0.7 mm long, 1.0—1.2 mm wide,
apically obtuse, densely pellucid punctate, the margin entire, densely glandular-
ciliate; pedicels cylindrical, 1.1-1.4 mm long, glabrous, epunctate. Pisti/-
late flowers unknown. Fruiting calyx cotyliform, coriaceous, 1.2—-1.4 mm long,
the tube 0.2—0.3 mm long, the lobes very widely ovate to oblate, 1.0—1.2
mm long, 1.1—1.3 mm wide, the apically acute, densely pellucid punctate,
abrous. Fruit globose, 0.8-1.2 cm

—

—

the margin irregular, opaque, entire, ¢
long and diameter, purplish at maturity, inconspicuously pellucid punctate.

Distribution.—Myrsine macrocarpa (Fig. 9) is endemic to the state of Amazonas,
Venezuela, growing at 1,500—2,000 m elevation.

130 SIDA 18(4)

Fic. 7. A. Habit, flowering branch of Myrsive macrocarpa Pipoly, drawn from R. Cowan GJ.
Wardack 31372 (NY isotype). B. Fruit of same, drawn from R. Cowan & J. Wurdack 31372
(NY isotype). C. Habit, lowering branch of Myrsine pellucida (Ruiz & Pav.) Spreng., drawn
from J. Luteyn & J. Pipoly 9370. D. Fruit of same, drawn from A. Gentry et al. 11090. FE.
Detail of branchlet apex and partial petiole with reddish glandular-papilae of same, drawn
from A. Gentry et al. 11090.

RICKETSON AND PIPOLy, Myrsine in Venezuela [131

Ecology and conservation status.—Myrsine macrocarpa is a riparian species, occurin g
at the margins of gallery forests in upland, wet savannas. Because its known
localities are so remote, and so wet, they are not particularly subject to dangers
from human intervention nor fire. Therefore, the species should not be considered
threatened.

Specimens examined. VENEZUELA. Amazonas: Cerro Hu: pera Rio Cunucunuma,
below Camp II, 1,000 m, 20 Dec 1950 (fr), B. Maguire et al. 29969 (NY-2 sheets); In vi-
cinity of Summit Camp, 1,800 m, 6 Dec 1950 (fr), B. Maguire et yi 30005 (NY); Along
right fork of Cafio de Dios in cumbre near Summit Camp, 13 Dec 1950 (fr), B. Maguire et
- 30180 (NY-2 sheets); yatamento Atures, lomas granfticas, Cafio Piedra, 115 km al

E de Puerto Ayacucho, 04° 54' N, 66° 54' W, 1,500 m, Sep 1989 (ster.), E. Sanoja et al.
on (MO).

Myrsine macrocarpa is most closely related to the vicariant M. pellucida
(Ruiz & Pav.) Spreng., but is easily separated by the prominulous second-
ary veins and pellucid punctations of the leaf blades, shorter pedicels, gla-
brous calyx lobe margins, and the large fruits.

LO. Myrsine pellucida (Ruiz & Pav.) Spreng., (Figs. 7C, ie 7E, 9), Syst.
Veg. 1: 64, 1825. Caballeria pellucida Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil.
1:280. 1798. Manglilla pellucida (Ruiz & Pav.) Roem. & Schult., ae Veg. 4:506.
1819. hee es (Rutz & Pav.) Mez in Engl., Pflanzenr. IV. 236(Heft 9):394.

1902. Ty J. HUANuco: Mufia, without elev., 1778-1788 (stam. fl), H. Ruiz
GJ. pa. an ain MA; Isorypes: F (NY Neg. # 12120), G-BOIS
ee ee Mez in Engl., Pflanzenr. IV. 236(Heft 9):395. 1902, syN. Nov. TYPE.
Y. PARAGUARI: Paraguari, without elev., Mar 1875 (fr), B. Balansa 2378
ee G; ISOTYPE: P).

Trees to 5(—10) m tall; trunk and branches orthotropic. Branchlets terete,
reddish glandular-papillose. Leaves spiral; blades coriaceous, elliptic, oblong,
oblanceolate or obovate, (6.5—)7.5—18 cm long, (2.7—)3.3—7.9 cm wide, apically
obtuse to subacute, basally acute, nitid above, densely reddish glandular-
papillate along the midrib above, the midrib impressed above, prominently
raised below, the secondary veins inconspicuous above and below, pellucid
or black punctate and punctate-lineate below, the margin flat, slightly de-
current on the petiole; petiole marginate, 0.5—1.0 cm long, densely red glandular-
papillate, glabrescent. Staminate inflorescence sessile, 8—12-flowered; floral bracts
coriaceous, ovate, ca. 1.3—1.5 mm long, 1.2—1.4 mm wide, apically broadly
rounded, long reddish glandular-ciliolate; pedicels obsolete to 1 mm long.
Staminate flowers chartaceous, 2-2.2 mm long; calyx cotyliform, ca. 1 mm
long, the cube 0.2—0.3 mm long, the lobes ovate, unequally divided, longer
than wide, 0.7—0.8 mm long. 0.5—0.7 mm wide, apically obtuse to sub-
acute, densely and prominently black punctate and punctate-lineate, the

margin alae glandular-ciliate, irregular, entire; corolla broadly eee

m long, the tube ca. 0.3 mm long, the lobes elliptic, 1.7—1.9 m

1132 Sipa 18(4)
long, 0.7-0.8 mm ube apically acute, densely and prominently black oes
and punctate-lineate, the margin entire, densely glandular-granulose throughout,
stamen 1.4—1.7 mm long, filaments not obvious, venation visible, ca. 0.3
mm long; anthers ovate to oblong, 1.5—1.7 mm long, apically apiculate,
the apiculum darkened, recurved ventrally, basally subcordate, the connec-
low, ca. | mm long, 0.5 mm diam.

—

tive black lineate; pistillode conic, ho
Pistillate inflorescence as in staminate. Pistillate flowers as in staminate but antherodes
|—1.1 mm long; pistil 2—2.3 mm long, |-1.3 mm diam., the ovary subglobose,
.| mm

—

|.2—1.3 mm long and in diam., the stigma carnose, prismatic, |—
long and in diam., the placenta ellipsoid, 1 mm long, the ovules 34, uniseriate.
Fruit globose, 3—3.5 mm diam., obscurely punctate and punctate-lineate.

Distribution.—Myrsine pellucida (Fig. 9) is known from the Venezuelan
coastal range in the state of Falcén, south and westward through the Andes
of Venezuela, Colombia, Ecuador, and Peru to Bolivia and adjacent Para-
guay, growing at (400—)1,000—2,600 m elevation. There are a few disjunct
populations in the western portion of the state of Bolivar, Venezuela, grow-
ing on diabasic intrusion areas in the tepuis.

Ecology and conservation status. —Myrsine pellucida is found along disturbed
roadsides through cloud forest and subpdramo life zones, and along mar-
gins of cloud forests. The disjunct populations in the state of Bolivar are on
diabasic intrusions in otherwise superimposed Roraima Sandstone formations.

Etymology. —The epithet ‘pellucida’ refers to the pellucid punctations of
the abaxial leaf blades and fruits.

sc imens examined. VENEZUELA. Barinas: oo. Pedraza, trail from Mesa de Canagua
1. 08° 34' N, 70° 37' W) to Pozo Negro (ca. 08° 32' N 70° 37' W), 400—600 m, 25 Nov
on (ft i L. Dorr et al, 7872 (MO, PORT); s 32 km NE of Altamira & 3 km NE of
Caldas, 08° 55'N, 70° 25' W, 1,000 m, 24 Jan 1984 (fr), J. Liuteyn GJ. Pipoly 9377 (BRIT,
O. NY, PORT). Bolivar: Distrito Cedefio, near Minerfa El Guaniamo, 06° 27' N, 65°
52' W, 300 m, May 1993 (ster.), EK. Diaz 1756 ergs near small Minerfa E1 Guaniamo,
06° 27' N, 65° 52' W, 300 m, May 1993 (ster), z 1759 (MO); Municipio Rat
Leoni, 64 km al SE de Pijiguaos, 06° 09' N, 66° 23' . 350 m, Jul 1989 (fl), Z ” Deleado
282 (MO). Falcén: Distrito Bolivar, mesa of Sierra de San Luis, ca. 1 km S of casero C Bates:
11° OS'N, 69° 42' W, 1,100 m, 31 Aug 1985 (fr), O. Huber & W. hi sbapiaase 1082 ae
MYE, MO, W):; Cucaire, Sierra de San Luis, 134 km post, 11° 08' N, 69° 46' W, 1,000 m,
20 Jun 1979 (fr), R. Liesner et al. 70601 (MO, VEN). Lara: Distrito Torres, ae Agua
Linda, near television cones towers ee él Fundo Orion, above the dei of Palmarito,
along sake Carrretera Lara-Zulia ca. 35 km E of El Venado, between Km 12-15, 10° LO'N,
0° 42' W, 6-7 Sep 1980 (fr), G. Biits i Stoddart 9745 (NY), Distrito Moran, trail
from Humocaro to Buenas Aires, Caserio, below Paramo Los Rosas, 09° 40! N, 70° 05! W,
2,600 m, 25 Jun 1979 (fr), R. ee et al. 7949 (MO); Distrito Moran, ca. 4.4 km W of
Humacaro Alto, 09° 36! N, 7( 'W, 1,400-1,500 m, 22 Jan 1984 (stam. fl), J. Luteyn
& J. Pipoly 9357 (MO, NY, aie . Mérida: ae without elev., 12 Nov 1952 (fr),
L. Bernardi 11 (NY): Antes de la Palmita, 700 m, 4 Feb 1955 (fr), L. Bernardi 1881 (NY,
VEN), Pueblos del Sur, 1,450 m, Jun 1955 (fr), L. Bernardi 2292 (NY 2-sheets); 26 km
NE of Mérida along road to Valencia, right bank of Chama River, 2,000 m, 18 Nov 1965

RICKETSON AND Pipory, Myrsine in Venezuela L153

(fl), F Breteler 3313 (NY); Mucuruba, Quebrada del pueblo, 2,700—2,800 m, 27 Jun 1930
(fl), H. Gehriger 270 (F, MO, NY); Distrito Rangel, E edge of town of Santo Domingo, 08°
52! N, 70° 43' W, 1,646 m, 24 Jan 1984 (fl), J. Lateyn & J. rae 9370 (E, MER, MO,
NY); Distrito Rivas Davila, paramo de La Negra, 23-30 km above (SSW) Bailadores on
road to La Grita, 2,500-2,900 m, 12 Apr 1984 . J. Luteyn Z M. Lebrén-Luteyn 9776
(NY); Along Hwy. 7, 21 km by road NE of Mérida and 9 km SW of Mucuruba, 1,900
29 ee 1979 (fr), M. Nee & M. Whalen 17016 (F); Anden, Umgebung von Timotes, ee
m, 22 Aug 1968 (fl, fr), B. Oberwinkler G Fo Oberw ee 12472 (M); Anden, Strafe Mérida,
Hee ner zwischen Tabay und Mucuruba, 2,000 m, 15 Dec 1968 (fl), B. Oberwinkler &
F. Oberwinkler 13909 (M); Vicinity of Timotes, ae m, 20 Jan 1928 (fl), H. Pittier 12652
Y); Disitrito Libertador, Caserfo El Chaba, entre Los Nevados y la poblacion de El
ey a orillas de la quebrada de El Hato, ays meridional de la Sierra Nevada, 2,400

, 14 Dec 1961 (A), L. Ruiz Terdn 824 (M QO); Disitrito Campo Elias, San José de
oe a Orillas de Quebrada de ee oe oo 2,400 m, 24 Mar 1967 (fl), L. Ruiz
oo et ies 3916 (MERE, NY). Tachira: Baas Zumbador, 14 km S of El Cobre, 2,500

. ur 1974 (fr), A. Gentry et al, 11090 (MO); Paramo Zumbador, 07° 58' N, 72° 05!
W, oe ren 4 Jan 1989 (fr), . Haln ee: (MO, NY); ea Quebrada
Cazadero, 16 km of San Cristobal, ( {'N, 72° 18' W, 650-900 m, 4 May 1981 (fl), R
Liesner & M. Guariglia 11812 a ae Laren 40 km S of La Grita along ioe to
San José de Bolivar, 07° 56'N, 71° 57! W, 1,560 m, 14 Apr 1984 (pist. fl, fr), J. Latyen G
M. Lebrén-Luteyn 9913 (MO, NY, VEN); Distrito Junin, Municipio Delicias, Caserfo Villa
Paez and vicinity, 1,900—2,300 m, 7 Mar 1962 (fr), L. Ruiz Terdn 935 (MERF, MO); Lower
slopes of Paramo de Tama to Paramo de Tama, 2,800 m, 20 May 1967 (A, fr), J. Steyermark
98562 (NY, VEN); Cabeceras del Rio ore ae steep slopes and cliffs of El Banco, belos
Cerro Las Copas, below Paéramo de Judfo, 20 kms al S de San Vicente de la Revancha, 35
kms S of Alquitrana, SW of Santa Ana, 2,500—2 70 )0 m, 15 Jan 1968 (fl, bud), /. He ie
et al. 100966 (NY 2-sheets); Sierra El Casadero, 13 km N of Rubio, between Las Dan
and Las Adjuntas, 07° 43' N, 72° 23' W, 900-1, a m, 12 Nov 1979 (fr), J. Steyer mate et
al, 120141 (MO). Trujillo: Distrito Carache, en Cuicas, La Guajira, 900 m, 25 Jun 1972
(fr), C. Benitez de Rojas 1459 (FB, MY).

Myrsine pellucida is a common plant throughout the Andes, particularly
easy to distinguish because of the sessile umbels and reddish glandular papillae
of the terminal bud and apical zone of the branchlets. This species, along
with Myrsine latifolia (Ruiz et Pav.) Spreng. have been misidentified as M.
guianensis, which does not occur in the Andes. Within Venezuela, it may be
confused with Myrsine macrocarpa, but is easily distinguished by the red glandular-
papillate branchles petioles and leaf midrib, smaller fruits and shorter peti-
oles. Another siginificant difference is that Myrszne pellucida is restricted to
soils on metamorphic rock, while M. macrocarpa is a tepui species.

11. Myrsine nitida (Mez) re (Figs. 6, 8A, 8B), Novon 1:210. 1991.
Rapanea nitida Mez, Repert. Spec. Nov. Regni Veg. 16:424. 1920. Typr. GUYANA
[VENEZU }. Me. oe 2,000 m, Dec 1909, (fl), E. Ule 8725 (HOLOTYPE: B-
destroyed, 1943; LecrorypE here designated K).

Rapanea roraimensis A.C. Sm., Bull. Torrey Bot. Club 67:296. ae SYN. Nov. ALyrsine
sealed C. Sm.) Pipoly, Nov yn 1:208. 1991. Type. VENEZUELA. Botivar: Mount
ma, SW slopes, 7,800 ft. [2,377 m], 11 Jan 1939 (fl), A. Palas 132 GIOLOTYPE:

a isoTypes: F, GH, MO, a US 2-sheets).

1134 Sipa 184)

wy WP

ail
i 8. A. Habit, flowering branch of ek nitida (Mez) Pipoly, drawn from A. Pinkus
32 (NY, holotype of Rapanea roraimensis A.C. Sm). B. Fruit of same, drawn from. J. Steyermark
7 ae 34 C. Habit, flowering branch of Myrsine eazanensis (Aubl.) Kuntze, drawn from J. Aublet

yn. (BM tsolectotype). D. Fruit ‘ same, drawn trom L. Delgado 215,

RICKETSON AND Pipoty, Myrsine in Venezuela iia b>)

Shrub or tree to 10 m tall; trunk and branches orthotropic. Branchlets ter-
ete, glabrous. Leaves spiral; blades coriaceous, elliptic or oblanceolate, (5.5)12—
17 cm long (2.7—)3.5—5.7(-7) cm wide, apically acute or rarely, obtuse, basally
acute to obtuse, slightly decurrent on the petiole, the secondary veins prominulous
above, obviously brochidodromous, the loop connections visible dull to somewhat
nitid above, conspicuously reddish punctate and punctate-lineate below, the
margin entire, flat; petioles marginate, 0.7—1.5 cm long, glabrous. Stami-
nate inflorescence fasciculate, S—9-flowered, the short shoots sessile to 3. mm
long; floral bracts ovate, 1—-1.2 mm long, 0.7—0.8 mm wide, apically acute,
densely and prominently black punctate-lineate; pedicels 1—1.3 mm long.
Staminate flowers S-merous, chartaceous, 3.4—3.6 mm long; calyx cotyliform,

2—1.4 mm long, the tube 0.2—0.3 mm long, the lobes ovate to deltate,

1-1.2 mm long, and wide, prominently black punctate-lineate medially,
apically acute, the margin entire, minutely glandular-ciliolate; corolla cam-
panulate, 3.4—3.6 mm long, the tube 0.7—0.9 mm long, the lobes oblong
to elliptic, 2.5—2.7 mm long, 1.2—1.3 mm wide, apically acute to obtuse,
densely and prominently black punctate and punctate-lineate, the margin
entire, glandular-granulose throughout; stamens 2.9—3.1 mm long; filaments
not obvious, ca. 0.7—0.9 mm long, the anthers ovate to oblong, 2—2.1 mm
long, 1.1—1.3 mm wide, apically apiculate, basally subcordate, the connectives
black punctate-lineate dorsally; pistillode conic, 2 mm long, 1 mm wide,
the sterile stigma conic, 4-lobed, hollow. Pisti/late inflorescence as in stami-
nate but pedicels 1-1.5 mm long. Pzisté//ate flowers as in staminate but 2.5—
2.9 mm long; calyx 1.5-1.7 mm long, the tube ca. 0.5 mm long, the lobes
ovate to delrate, 0.9-1 mm long and wide, apically acute; corel 1.8-2
mm long, the tube 0.7—0.8 mm long, the lobes elliptic, 1.1—1.3 mm long,
ca. 0.5 mm wide; staminodes ca. 1.5 mm long, the filaments not obvious
but venation visible, ca. 0.7—0.8 mm long, the antherodes malformed, 0.5—
0.8 mm long, ca. 0.5 mm wide, apically acute, basally subsagittate; pistil
2.5—2.7 mm long, the ovary globose, 1.1—1.3 mm long and diam., the stigma
lacinate, 4-lobed, translucent, carnose, 1.5—1.7 mm long, 1—1.2 mm diam,
the placenta ellipsoid; ovules 3—4, uniseriate. Friit aisha s 3.5 mm long
and in diam., obscurely pellucid punctate.

Distribution.—Myrsine nitida (Fig. 6) is known from the state of Bolivar in
Venezuela, and adjacent Guyana, growing at (830—)1,800—2,450 m elevation.

Ecology and conservation status. —Myrsine nitida occurs in rocky knolls in
savannas, along the margins of premontane forests, C/asia-Magnolia ripar-
ian forests, and in Mora forests. Because these forests produce more timber
than any of those surrounding them, this species should be considered threatened.

Etymology.—The specific epithet refers to the shiny nature of the adaxial
leaf surface.

1136 Stpa 18(4)

Specimens ees GUYANA. Cuyunt-Mazaruni: Pakaraima Mts., NE plateau of
Ayanganna, 05° 23' 59° S8' W, | ees 650 m, 3 Nov 1992 (ster.), B. Hoffman 3230
(BRIT, US). seen Bolivar: 0-1 km NE of El Paujf, 04° 30'N, 61° 35' W, 830—
900 m, 5 Nov 1985 (fl), R. Liesner oe (MO, NY, VEN): Gran Sabana, lt tepui, W-
facing slopes below low escarpment, 7,000—8,000 ft [2,134—2,438 m], 17 Mar 1952 (fl),
B. Maguire 334068 (BRIT, NY), slopes below upper most W-facing escarpment, 21 Mar

1952 (fr), 33497 (NY); Mount Roraima, ae facing slopes between Rond6én cei and base
yf bluffs, 2,040—2 ee m, 30 Sep 1944 (fr), J. Steyermark 58948 (F, NY); Prari-ceput, along
be ise of E-facing bluff, 2,410—2,450 m, 7 ee 944 (EH), J. Steyermark a (F, a Sororopan-
tepul, crest of cerro between E & W end, 2,255 m, 14 Nov 1944 (fl), J. Steyermark 60108
(F, NY, US); Chimanta Massif, above SE-facing upper shoulder on re leading tosummit
unre tepui, 2,200-2, 300 ym, 20 Jun 1953 (fl), J. Steyermark 75834 (EF, MO, NY); Chimanca
f, Agparaman-tepui, SE- facing forested slopes below escarpment, |,880—1,955 n
. Wares G Fi 1160 (N

Myrsine nitida appears to be most closely related to M. guianensis, but is
separated by the conspicuously reddish punctate-lineate abaxial leaf sur-
face, the acute or rarely, obtuse leaf apices and habitat.

12, oe ee (Aubl.) Kuntze, os 8C, 8D, 9), Revis. Gen.
= 402. 1891. Rapanea guianensis Aubl., . Pl. Guiane. Fr. 1:121, ¢. 46.
35, ae: FRENCH GUIANA. Without ee aes elev., without date (fl);
J. Aublet sn. Lecrorye: P, here designated (BRIT & MO Neg.); IsoLecroryPE: BM
(MO Neg. # 4321)).

Rapanea ee Pohl ex Miq. in Mart., Fl. Bras. 10:308. 1856. Type. BRAZIL. With-
out locality, without elev., without ae (A, fr), Pohl 4384, 6077 (syNrypEs W; photo
SI). We c
and their duplicates can be assembled.

Myrsine ovalifolia Mig. in Marc., Fl. Bras. 10:313. 1856. Rapanea ovalifolia (Miq.) Mez
in Engl., Pflanzenr. 1V. 236( Tele 9):391. 1902. Type. BRAZIL. Alagoas, without elev.,
Jun 1838 ( fl), Gardner 1350 (HoLorypE: G; IsoryPE: BM)

OP agg

efer leaee aa of this binomial uncil all material cited in the protologue

Trees to O(—-15) m tall; trunk and branches orthotropic. Branchlets terete,

glabrous, the bark of lower branches thick and corky. Leaver spiral; blades
coriaceous, elliptic, oblong, oblanceolate or obovate, 7-18 cm long, (2.7-
)3—7.5 cm wide, apically obtuse to broadly rounded, basally acute, the midrib
impressed above, prominently raised below, the secondary veins prominulous
above, penninerved, not obviously brochidodromous, nitid above, pallid below,
inconspicuously black punctate or black punctate-lineate below, the mar-
gin entire, flat, somewhat revolute basally; petioles marginate, 0.5—1.5 cm
long, glabrous. Staminate inflorescence fasciculate, S—8-flowered, on short
perrennating shoots accrescent by accumulating girdling floral bracts; floral
bracts chartaceous, oblate, 0.5—0.6 mm long, 0.9—1.0 mm wide, apically

—

broadly rounded, the margin entire, densely long glandular-ciliate; pedicels
1.2—2 mm long. Staminate flowers S-merous, chartaceous, hyaline, 3.5—3.8 mm

mm long, the tube ca. 0.3 mm long, the

long; calyx campanulate, 1—-1.2
.7—0.9 mm long and wide, apically narrowly

lobes widely ovate to deltate, (

—

RICKETSON AND Pipoty, Myrsine in Venezuela 1B oi

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Fic. 9. Distribution of Myrsine pellucida (#), M. guianensis (QO), M. macrocarpa (g), M. perpauciflora
(~) and M. minima (A) in Venezuela and bordering Guyana region.

L138 Stipa 18(4)

acute to attenuate, prominently black punctate and punctate-lineate medi-
ally, the margin entire, glabrous; corolla campanulate, 3.5—3.8 mm long,
the tube 0.7—0.9 mm long, the lobes elliptic to lanceolate, 2.7—3 mm long,
0.9—l1 mm wide, apically sharply acute to attenuate, densely and promi-
nently black punctate and Se the margin entire, glandular-
granulose throughout; stamens 2.2—2.5 mm long, the filaments not obvi-
ous (but veins visible) 0.7—0.9 mm loan the anthers ovate, 1.5—1.7 mm
long, 0.7—0.8 mm wide, apically apiculate, the apically curved inward, basally
subcordate, the connective with one prominent black punctation at point
of attachment; pistillode conic, 1.3—1.5 mm long, 0.7—0.8 mm diam., hol-
low or bearing a sterile placenta. Pisti//ate inflorescence as in staminate but on
perrenating shoots to 6 mm long; pedicels |.5—2 mm long. Pésti/late flowers
as in staminate but calyx 1.1—1.8, the tube ca. 0.2 mm the lobes ovate,
1.1—1.3 (1.5) mm long, 0.7—0.9 mm wide; corolla 2.5—3.5 mm long; the
tube as in staminate, the lobes 2.5—3 mm long, 0.8—1.2 mm wide; staminodes
resembling stamens except antherodes broadly ovate 1.0—1.3 mm long, 0.6—
0.8 mm wide; pistil conic, 1.5—2.5 mm long,O.8—1 mm diam, the stigma
conical, with 4 twisted lobes, the placenta conical, the ovules 3—5, uniseriate.
Fruit globose, 3.5—5 mm long and in diam., prominently black punctate.

Distribution.—Myrsine guianensis is known from French Guiana and ad-
jacent Brazil westward through Suriname and Guyana to Venezuela (Fig.
9), growing at (100—)400—1,400(—2,800) m elevation. Reports of this spe-
cies from Colombia, Ecuador, Dena Bolivia and Paraguay are based on
misidentifications. In Venezuela, it is found throughout the states of Bolivar,
around the margins of Amazonas, in Delta Amacuro, Monagas, Sucre, Nueva
Esparta, Anzoategui, Miranda, D.F., Aragua, and Falc6n growing in sandy,

—

savanna-like situations.

Ecology and conservation status, —In Venezuela, Myrsine guianensis is known
from primary and secondary lowland riparian forests on reddish or quartz-
ite sands, premontane forest, cloud forests and rarely, upland savannas.

Etymology —The epithet refers to the species’ type locality, French Guiana.

Specimens examined. GUYANA. Mazaruni-Potaro: Upper Potaro River region, sum-
mit of Mt. Wokomung, 05° 05' N, 59° 50' W, 9 Jul 1989 (fr), B. Boom & G. Samuels 9138
(MO, NY); Upper Potaro River region, aes es -Orinduik, ca. 1.5 hr walk from
Koponang, 04° 57' N, 59° 53' W, 790 2 Jul 1989 (fr), B. pe . G. Samuels 9285
(MO, NY): Kaieteur Plateau, forest ae eal from Plane-landing to Kaieteur Falls, ca.
1,400 m, 11 Feb 1962 (fl), R. Cowan & T. Soderstrom 1826 (K, US); E Berbice- Corentyne
pe ees 3 Savanna; Canje River, 1O km S$ of Mora, 185 en S or mouth of Canje,

33'N, 5S YW, 10-20 m, 29 Oct nen H), L. Gillespie et al, 2510 (US); Potaro-
chi oe at southern base of Mt. Kopinang along trail from Kopinang to Orinduik,
05° 00' N, 59° 55' W, 500- 600 m, ae cae W. Hahn et al. 4435 (MO, US); Potaro-
Siparunit Region, Kato, 04° 40' N, 59° 55' W, 750 m, 11 Mar 1989 (fr), W. Hahn et al.
5603 (MO, US); E. Berbice- . ore ey gion: W bank of Canje Rive, Cow Savanna, c
km N of Digitima Creek, 1— , 14 Apr 1987 (fr), J. Pipoly et af. 11582 (FDG, MO, <<

RICKETSON AND Pipoty, Myrsine in Venezuela 1139

US)Upper Mazaruni River basin, Mc. Ayanganna, on shoulder of E flank, ca. Thompson
camp, 1,418 m, 12 Aug 1960 (fr), S. Ti/lert et al. is soe sheets); Upper Mazaruni

River basin, Kukui River, in Mora forest bordering r at Adaro river mouth, ca. 300 m,
5 Sep 1960 (fr), S. Tillett & C. Tillett 45290 (NY-2 eee US). VENEZUELA. Amazonas:
Beyond soccer field, San Carlos de Rio Negro, 115 m, 17 Sep 1975 (fr), P. Berry 1367 (MO);
San Carlos de Rio ees ca. 20 km S of confluence of Rio Negro and Brazo Casiquiare,
near radar site, 3.4 km NE of San Carlos on Solano road, 01° 56! N, 67° 03' W, 119 m, 13
Mar 1979 (fr), H, Clark oy (MO, NY); Serrania de Tapirapecé, Campamento Tamacuari,
trail between 2 camps, 01° 14'N, 64° 40! W, 1,400 m, 10 Feb 1989 (fr), A. Henderson et al.
1022 (BRIT, NY); Department Atures, cumbre del Cerro Yavi, en las cabeceras del Rio
Parucito, afluente oriental del Rio Manapiare, en el sector NE de la cumbre, 05° 43'N,
65° 52! W, 2,100 m, 29 Oct 1986 (fr), O. Huber 11853 (NY); Department Atures, cumbre
del Cerro Coro-Coro, en las Peer e nor-occidentales del Rio Manapiare, sector NW de la
Serrania Yutajé, 05° 46' N, 66° 1 W, 2,200 m, 12 Nov 1987 (fr), O. Huber 12318 (US);
Sierra Parima, Vecindades de sea hi, Rio Maid uni, a unos 6—7 km al O de la frontera
Venezolana-Brasilera, 03° 49' N, 64° 36' O, 795-830 m, 18 Abr—23 May 1973 (fl), J.
Steyermark 106968 (MO, NY); 107113 (F, MO, NY). Anzoategut: Distrito Bolivar, just S
of E harper Fila El Purgatorio, 9 airline km NE of Bergantin, 10° 02' N, 64° 17! W,
1,100 m, 24 Nov 1981 (fr), G. Davidse & A. Gonzdlez 19400 (MO, NY, VEN), 19402
cs. Aragua: Maracay, without elev., 1934 (fl, fr), P. Vogel 1118 (M); Arriba de Guamitas,
.N. Aragua, 780 m, 15 Jun 1939 aan Es ee 11125 (& MO). Bolivar: Near summit

Qu

of Cerro anal ironstone area, Ciudad Piar, 750 m, 10 Apr 1954 (fl), L. Arésteguieta 2178
(NY 2-sheets); Disitrio Raul Leoni, Rio Ariza, 55 a NE of San Se de la Paragua,
O79 1 _ 64° 13' W, 485 m, Jun 1989 (fr), L. Delgado 215 (MO, NY); Disitrito Gran

Sabana, Zona Minera El Polaco, 04° 32' N, 61° 26' W, 950 m, i o (ster.), W. Diaz y
S. Elcoro : 280 (MO); Disitrito ue Leonf, Bajo Caronf, Sector III, Cerro Altamira, 07° 27!
N, 63° 13' W, 300 m, May 1994 (fl), W Diaz 2515 (MO); Municipio Piar, 3 km NE of
Hato ie Nek 07° 28'N, a 36! W, 280 m, May 1986 (fr), A. Ferndndez 2946 (US);
Distrito Piar, Serranfa Quiribay, 07° 49'N, 62° 43' W, 280 m, 26 Jul 1989 (fr), F Flores &
E. Pérez 557 (MO); Guyana, villa de Upata, without elev., 1864 (fl), D. de Grosourdy 13 (P);
Distrito Bene Meseta del See S-sloping plain near edge of NE section of the
meseta, near the summit, 05° N, 63° 29' W, 1,400 m, 27 Mar 1985 (fr), O. Huber
10389 (MYF, MO, NY); ae aoe Meseta del Guaiquinima, SE sector of the sum-
mit, 05° 51' N, 63° 25' W, 1,350 m, 26 Sep 1985 (fr), O. Huber & G. Medina C. 10916
(MYE, — Distrito Cedefio, Se rrania Guanay, NE sector, sloping toward the S and SW, at
the easternmost headwaters of Rfo Parguaza, 05° 55' N, 66° 23' W, 1,700 m, 20-28 Oct
1985 (fl bud), O. Huber 11062 (MYE, NY); Distrito Piar, Macizo del Chimanta, W sector,
SW range of Apacard-tepui, near the connection with Abacapd-tepui, at NW headwaters
of Rio Tirica, 05° 17! N, 62° 16' W, 2,100 m, 8-10 Mar 1986 (fr), O. Huber 11456 (BRIT,
MYE, MO, NY 2 sheets); Disitrito Roscio, savannas at W fools: and SW summit of the
Aldisbigice de Nuria, ca. 5O km N of Tumeremo, 07° 37' N, 61° 37' W, 300-450 m, 7
May 1986 (fl buds), 0. Haber 11640 (MYF, NY); Distrito Heres, Mest del ee
iy sloping a ae the SE border of the meseta, at W headwaters of Rio Aberaim
5° 47' N, 63° 48' W, 1,400 m, 20 May 1987 (fl), O. Huber & V. Rull 12273 (MYF, a.
oo ene cn Meseta de Jaua, headwaters of Rio Marajano, tributary of Rio Cacaro,
04° 48'N, 64° 32' W, 1,750-1,800 m, 20 Nov 1989 (fr), O. ees 13005 (MYE, MO);
Uaipan-teput, between the W & E Peaks of Uaipan, ca. 1,500 m, 4 Mar 1967 (fr), 7) Koyama
& G. Agostini 7474 (NY, VEN); Cerro Guaiquinima, Rio i. laterial S drainage of
“North Valley”, 1,700 m, 2 Jan 1952 (fl), B. Maguire 32917 (NY); Ptari-tepui, NW slopes,
1,500—2,000 m, 17 Dec 1952 (FR), B. Maguire & J. Wurdack 33896 (NY), Ptari-teput, in

1140 Stipa 18(4)

vicinity of “Cave Rock” camp, below S face of mountain, |,600—2,000 m, 14-19 Aug 1970
(bud), H. Moore et af. 9738 (BF, NY, US); Ptari- tepui, S slope, es m, 24 ae 1983 (fr), 7.
Pipoly 0399 (NY, VEN), Prari-tepui, S-facing slopes, just NE of “Cave Rock”, 1,800—1,850
m, 4 Nov 1944 (fr), J. Steyermark 59808 (F, NY); Ptari-tepui, at base of cerro along Rio
Karuai, 1,220 m, 27 Nov 1944 (ster.), J. Steyermark 60621 (F, MO); Chimanta Massif, NW
part of summit of Abdcapa-tepui, 2,125—2,300 m, 13 Apr 1953 (fr), J. Steyermark 74954
(F, MO, NY); Chimanta Massif, on plateau of SE-facing upper shoulder of Apacara-tepui,
2,000 m, 19 Jun 1953 (fr), J. Steyermark 75738 (F, a Y Chimanta Massif, below
Upper Falls of Rio Tirica above Summit Camp, 1,940 m, 7 Feb 1955 (fr), J. Steyermark &
J. Wurdack 594 (BRIT, F, MO, NY); Altiplanicie de Nuria, vicinity of campamento Nuria,
NE of Hato de Nuria, 530 m, 17 Jul 1960 (ster.), J. Steyermark 86438 (NY); Altiplanicie
de Nuria, vicinity of camp, 5 kms from Hato de Nuria, E of Miamo, 400 m, 12 Jan 1961
(fr), J. Steyermark 88366 (NY); along Rio Churtin at ne of “Second Wall” of sandstone, N
from camp, 5 km to the NE, 1,660 m, 12 May 1964 (fl), J. Steyermark 93748 (NY); Distrito
Heres, Cerro ane 1,420 m, 13 Jan 1981 (Al ae ), J. Steyermark et al. 124036 (NY);
Distrito Piar, Cerro Torribio, Bajo Caronf, Sector II, 07° 04' N, 62° 05' W, 100 m, May
1994 (fr), A. Valera 563 (MO). Delta Amacuro: De partamento Antonio Diaz, vicinity
Cano Jotajana, tributary of a ei age NW of Epafia near boundary with oar
Peng 09° 15'N, 61° 10! W, 50 m, 20 Oct 1977 (ster.), J. Steyermark et al. 115090
(MO). Distrito ats Bae Pes ia: elev., Dec 1924 (fl), A. A//art ee 3 a
Parque Nacional “El Avila,” near Los Venados, 1,650 m, 22 Dec 1975 (fl), L. _ &
O. Huber 300 (NY); Reforested hills of the Caracas Botanical Gardens, 870-980 m 1 Jul
1974 (fr), P. Berry 213 (MO); Calvario de Caracas, without elev., without date (fi), Ernsi
1088 (BM); Prope coloniam Tovar, without elev., 1854-1855 (fl), A. Fendler 760 (MO,
NY); Parque Nacional El Avila, above Naiguatd, along the right of way for Electricidad de
Caracas between the stream of the Rio Camuri and the stream of the Rfo Mazares en selva,
10° 34.5' N, 66° 42.3' O, 800 m, 11-12 Apr 1992 (scer.), W. Meizer et al, 2069 (BRIT);
Jardin Botanico, Caracas, reforested slopes facing N, without elev., 14 Jun 1960 (fr), /.
Steyermark 86309 (NY). Caracas, without elev., 14 Jan 1892 (A), J. ae 560 (C 2-sheets).
Falcén: Peninsula Paraguand, Cerro Santa Ana, 650 m, 15 Dec 1964 (fl), F. Breteler 4292
(MO, NY); 4299 (EF MO, NY): Peninsula Paraguana, Cerro Santa Ana, 650 m, Dec 1953
(A), 7 Lasser & L. Aristeguieta 3410 (F 2-sheets, VEN); Cerro Santa Ana, ascension del lado
S desde el pueblo de Santa Ana, 750 m, 24 Jan 1966 (fl), J. Steyermark & A. Braun 94642
(NY); Penfnsula ee ate Cerro Santa Ana, without elev., Jan 1939 (A, fr), F Tamayo
797 (BF). Miranda: Carretera de Tacata-Loma del Hierro sabana antropica, without elev., |
Jan 1988 (fl), N. Renin 2477 (MO, NY). Monagas: San Antonio-Cumané Road, without
elev., 24 Oct 1948 (A, fr), B. Maguire et al, 27251 (NY); Cerro Negro, above La Sabana de
las Piedras, NW of Caripe, 1,500—2,180 m, 15 Apr 1945 (Al),J. Steyermark 62075 (BF). Nueva
plone Isla de eae pe Catia Copey, 500-900 m, Jul 1984 (11), FE Delascio & A. ey
12223 (MO); (fr), 7 5 (MO); Copey, 900 m, Dec 1951 (A), H. Gines 2815 (US), -
(US); Is la de Margarita, e erro Copey, S of Santa Ana, at and near summit, 11° 02— 03)
63° 55' W, 800-850 m, 24 Mar 1985 (fr), /. echt al, 131074 (MO). Sucre: Valley
of Cocollar, 820 m, 28 ne 1945 (fr), J. Steyermark 62377 (F); Cerro Turumuquire, N-fac-
ing slopes above La Trinidad, SW of Cocollar, 2,100—2,2 Ht m, 5 May 1945 (fr), J. Steyermark
62560 (F); Distrito Sucre, La Lomita, between La Sabana and Zurita, 500-600 m, 18 Aug
1973 (A), J. Steyermark et al. 107789 (MO, NY). Wrruour LocaLiry: Coticita, without
elev., 17 Jun 1917 (fl, bud), H. Curran & M. Haman 1069 (NY)

pee

Myrsine gitanensis (Aubl.) Kuntze has long been confused with M. pe/lu-
cida and M. latifolia (Ruiz & Pav.) Spreng., both Andean species. In the Caribbean

RICKETSON AND Pipoty, Myrsine in Venezuela 1141

and Central America, including the coast of the state of Delta Amacuro, it
has been confused with Myrsine cubana A. DC. (often reported as M. floridana
A. DC., a nomen superfluum, or M. punctatum (Lam.) Stearn, non (H. Lév.)
Wilbur). In Brazil, it has been confused with several other species, includ-
ing Myrsine monticola Mart. (Harvey & Pipoly 1995) and M. wmbellata Mart.
In Venezuela, Myrszne guianensis is most easily confused with M. nitida, from
which it is easily distinguished by the obtuse to broadly rounded leaf api-
ces, with inconspicuously black punctate and punctate-lineate abaxial sur-
faces, the ovate calyx lobes, larger, black punctate fruits and the cloud for-
est to lowland savanna habitats.

DOUBTFUL AND EXCLUDED TAXA

Myrsine schomburgkiana Mig. in Mart., Fl. Bras. 10:315. 1856.= ee
schomburgkiana (Miquel) Baill., Hist. Pl. 11 293, 1892, (Sapotaceae). Tyre. GUYANA.
ithout further locality, without elev., iehout date, R. Schomburgk 968 — PE:

U; tsoTyPe: BM). See T. Pennington, Fl. Neotrop. 52: 240. 1990.
ee a Gleason, Bull. Torrey Bot. Club 58:447. 1931. Type. VEN-
AMAZONAS: Summit of Mount Duida, 4,800 ft [1,463 m], 5 Jan 1929

Te e oe 594 (HOLOTYPE: NY

This taxon is actually a species of Micropholis (Sapotaceae), and the type
was annotated “Micropholis sp.” by Pennington in 1985. However, disposition
of this name was not made in his Sapotaceae monograph (Pennington 1990).

ACKNOWLEDGMENTS

We thank the Missouri Botanical Garden and the Flora of the Venezu-
elan Guayana Project, for funding that allowed J. Ricketson (MO) to travel
to Fort Worth, and for Pipoly to make visits to MO. We also thank the
curators of the herbaria cited for loans of specimens. We are also grateful to
those who have been so instrumental in assisting us in our work, including
Gerrit and Jeany Davidse, Linda Oestry, Mary Bard, and Catherine Mayo,
(MO), Barney Lipscomb, and Jim Rivers (BRIT).

REFERENCES

ANDERBERG, A. and B. STAHL 1995. Phylogenetic interrelationships in the order Primulales,
with special emphasis on the family circumscriptions. Canad. J. Bot. 73:1699—1730
C. CHEN and J. Pipory. 1996. Myrsinaceae. In: Wu Zheng-yi and P. Raven, or pe of
China. ee Press, Beijing and Missouri Botanical Garden, St. Louis, MO. —38.

CULLEN, J. 1978. A preliminary survey of ptixis in the Angiosperms. Notes ae Bot.
Gard. aan 37:161-214

Hate, F., R.A. OLDEMAN, and P.B. TOMLINSON. 1978. Tropical trees and forests. An archi-
tectural analysis. Springer-Verlag. New Yor

Harvey, Y. and J. Pipory. 1995. Myrsinaceae. Ta) B. Stannard, ed. Flora of the Pico das
Almas, we pada Diamantina, Bahia, Brazil. Royal Botanic Gardens, Kew, Richmond,
U.K. Pp. 487-491.

1142 Stipa 18(4)

HIcKEY, 1984. A revised classification of the architecture of guia leaves. In:
CR. see and L. Chalk, eds. Anatomy of the dicotyledons. Vol. 1. Systematic anatomy
of leaf and stem, with a brief history of the subject. Clarendon Oxtord, U.K. Pp.
25-39.

Huser, O. 1995. Vegetation. In: J. Steyermark, P. Berry, and B. Holst, eds. Flora of the
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St. Louis. Pp. 97-160.

LINDLEY, a 1848. Iluserated dictionary of botanical terms. Excerpt from illustrated dictio-

—

nary of botanical terms by John Lindley. [Reprint, With an Introduction by Alice Eastwood,

Stanford University, School of Earth Sciences, 34]

Metcare, C.R. 1984. Some basic types of cells and tissues. In: C.R. Metcalfe and L. Chalk,
eds. Anatomy of the dicotyledons. Vol. 1. Systematic anatomy of leaf and stem, With a
Brief History of the Subject. Clarendon Press. Oxtord, U.K. Pp. 54-62

Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9): 1-437.

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a}

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2

Pipoiy, J. 1983 a ee a wont of Cybianthus (Myrsinaceae): III. A
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99 L. Systematic studies in ne genus Myrszve L. (Myrsinaceae) in Guayana
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l¢

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Publ. London. 4:7 1—

RICKETSON AND Pipoty, Myrsine in Venezuela 1143

NUMERICAL LIST OF MYRSINE TAXA

L. Alyrsine dependens 7. Myrsine resinosa
2a. Myrsine coriacea subsp. coriacea 8. Myrsine picturata
2b. Myrsine coriacea subsp. reticulata 9. Myrsine macrocarpa

3. Myrsine Maguirec ana 10. Myrsine ca ida

4. Myrsine minima Ll. Myrszne at

5. Myrsine andina 12. Myrsine uae

6. Myrsine perpanciflora

LIST OF EXSICCATAE
Figures in parentheses refer to the numbers from the numerical list of
taxa. Collection numbers in boldface type indicate type specimens.

Agostini, G. 174 (2a); Allart, A. 513 (12); Alston, A. 5515 (2a); 6172 (2a); Anderson, W. 13461
(2a); Arisceguieta, L. 2178 (12); 3606 (1); 5498 (2a); Aristeguieta, L. & O. Huber 300 (12) Oe
J.s.n. (12); Aymard, G. et al. 5190

Badillo, V. 7339 (2a); Baruch, Z. 95 (2a); Bernardi, L. 11 (10); 214 (1); 1881 (10); 2292 (10): Bernardi,
L. etal. 17136(1); se 1): pee. P. 213 (12); 1367 (12); Boom, na eine ee 9285
2); Boom, B. et al. 5 (3); Breteler, EF. 8 (2a); 3242 (2a); 3313 (10); 3635 (2a); 4289 (2a);
4292 (12); 4580 (2a); ee er, J. 4629 (1); Bruijn, J. de 988 (2a); ace ee M. Fucci 6050 (2a);
Bunting, G. & A. Stoddart 9745 (10).

—

oan A. & H. Meneses 1421 (2a); Cardozo, A. & H. Rodriguez 1218 (2a); Cardozo, A. €
(2a); 1344 (2a); Chapin, A. & F. Jacquemoud 13136 (2a); 13296 (2a); Clark, H. 7094 (12); as
S.& si Diaz ue 2435 (1); Clemants, S. & J. Dugarre 2411 (1); Colella, M. & O. Huber 451 (2a); 465

(2b); Cowan , R. oS Soderstrom 1826 (12); Cowan, R. & J. Wurdack 31372 (9); Croat, T. 59551
(3); Aten J. etal. 28185 (1); ven N. 931 (2a); 1107 (2a); 1256 (2a); 1335 (2a); 1348 (2a);
Curran, H. 2114 (2a); Curran, H. . Haman 1069 (12).

ee: G. & A. Gonzalez agree Ges ee 19540 (2a); 19606 (2a); Davidse, G. et al. 20813

2a), Delascio, F. & A. Gonzalez 12223 (12); 12255 (12); Delgado, L. 215 (12); 282 (10); Diaz, E.
Hee (10); Diaz, W. 1759 (10); 2515 os ae . & S. Elcoro 1580 (12); Dorr, L. & L. Barnett
S157 (2a); 3162 (2a); 5602 (2a); 7598 (2a); Dorr, L. et al. 5693 (2a); 7872 (10); Dumont, K. er al.

Elcoro, S. & N. Vera 225 (2a); Ernst, A. 1088 (12).

Farney, C. et al. 905 (4); Fendler, A. 758 (2a); 760 (12); Fernandez, A. 2946 (12): Flores, F. & E. Pérez
557 (12); Funck, H. 481 (1); 550 (1); Funck, H. & L. Schlim 58 (1); 140 (2a).

Gehringer, H. 28 (1); 270 (10); 401 (2a); Gentry, A. et al. 11090 (10); Gil lespie, L. et al. 2510 (12);
Gines, H. 2815 (12); 3461 (12); Grosourdy, D. 13 (12); Guevara, L. C. de 3261 (2a).

Hahn, W. & F. Grifo 3460 (2a); 4971 (5); 4983a (10); Hahn, W. et al. 4435 (1.2); 5603 (12); Henderson,
A. et al. 1022 (12); Hernandez, L. & N. Dezzeo 114 (2a); Hoffman, B. 3230 (11); Huber, O. 9269
(4); 10389 (12); 11062 (12); 11456 (12); 11593 (4); 11640 (12); 11698 (4): l pieces 12318 (12);
13005 (12), 13071 (3); Huber, O. & C. Alarcén 7375 (2a); 7931 (2a); Huber, O. & N. Deno 8590
4); Huber, O. & L. Izquierdo 12802 (2b); Huber, O. & G. Medina C. 10916 (12); Huber W.
Morawetz 10 823 (10); Huber, O. & V. Rull 12273 (12); Huber, O. et al. 8786 (2a); 10100 (4): ees
& A. Bonpland s.n. (1).

Karsten, G. s.n. (2a); Koyama, T. & G. Agostini 7474 (12); Kral, R. & A. Gonzalez 70455 (2a).

Lasser, T. & L. aye 3410 eae 3418 (2a); Licata, cons er 1. 650 (2a); Liesner, R. 19543 (11);
23107 (4); 23210 (A); 23296 (4); 23426 (4); 23465 (7); 235 re. Liesner, R. & A. Gonzalez 9933
(2a); 10713 (2a); ie R. & M. Guariglia 11655 (5); ee (10); Liesner, R. et al. 7661 (10); 7949
(10); 8202 (2a), 8232 (2a); 8389 (2a); Linden, J. 108 (5); 958 (1); Luceyn, J. & M. Lebrén-Luteyn

1144 Stipa 18(4)

9776 (10); 9913 (10); ; Luteyn, J. & J. Pipoly 9280 (
9377 (10); on J. et al. 9386 (2a); 9387 (2:

Maguire, B. 32917 (12); 33465 ioe 33468 (11); 33497 (11); 33535 (8); Maguire, B. & J. Wurdack
33896 (12); sea B. ec al. * eae Era 30005 (9); 30180 (9); 42090 (8); 42318 (3);
42375 (8); 42376 (8); 42393 Or 2403 (3), 60450 (4); Manara, B. s.n ); Marcano-Berti, L. 804
(1); Meier, W. 14 (2a); 1097 (1); 13 i; (2a); 1414 (2a); 1603 (2a); Meter, Ww. & G. Bronner 3523 (2a);
Meier, W. & O. Carrero 869 (2a); cae ~etal. 2069 (12); Moore, H. et al. 9738 (12); Morillo, G.
592 (2a); 2626 (2a); Morillo, G. et al. 2887 (7); 9606 (2a); Moritz, J. 153 (2a); 360 (2a); 1175 (2a).

Nee, M. 30697 (6); Nee, M. & M. Whalen 17016 (10).
Oberwinkler, B. & E Oberwinkler 12472 (10); 13909 (10).

Pinkus, A. 84 ie 132 (11); Pipoly, J. 6399 (1 2); 6450 (2a); Pipoly, J. & G. Aymard 6566 (1); Pipoly,
— et a ave. - 6486 (2a); 6512 (1); 7141 (4); 11582 (12); Pittier, H. 12652 (10); 12793 (2a);
2a).

2a); 9328 (2a); 9335 (2a); 9357 (10); 9370 (10);

~

Ramirez, N. 796 (2a); 2084 (2a); 2477 (12); Ramirez, N. et al. 4452 (2b); 4666 (2a); Renner, S. 2096
(6); Rojas, C. de & F. Rojas 3682 (2a); Rojas, C 7 Dente de 1459 (10); Ruiz pee L. 824 a 935
(10); 1297 (2a); 1436 (2a); 1669 (1); Ruiz Teran, L. & M. Lopez-Figueiras 126 (2a); Rufz Teran, L.

1. 3916 (10).

Sanoja, E. et al. 2988 (9); Sobel, G. & J. Strudwick 2151 (2a); Sobel, G. et al. 2021 (2a); Stein, B. &
A. Gentry 1525 (6); Steyermark, J. 55022 (2a); 55646, (1); 56588 (2a); 57205 (5); 58848 (4); 58948
(11); 58983 (2b); 59808 (1 2); 99826 ( (2a); 59948 (11); GOLOS (11); 60621 (12); 60726 (7); 60737
(2a); ela 8 (2a); 61355 (2a); 61445 (2a); 61615 (2a); 61927 (2a); 62075 (12); 62377 (12); 62560
12): 62721 (2a); 63007 (1); 74954 (12); 75738 (12); 75834 (11); 75931 (4); 86192 (2a); 86309
(12); eects (12); 88366 (12); a 18 (12); aa (2a); 97985 (4); 98431 (2a); 98562 (10); 99182
(2a); 100998 (5); 104785 (1); 106283 (2a); 106968 (12); 107113 (12); Steyermark, J. & A. Braun
94642 (12); Steyermark, J. & J. Wurdack 594 (12); 806 (2b); 843 (4); 1194 (2b); Steyern mark, J. etal.
oo \); 107789 (12); 115090 (12); 120141 (10); 121737 (2a); 124036 (12); 124328 (4); 128268
(4); 131074 (12).

Tamayo, F. 436 (2a); 797 (12); Tanner, E. & V. Kapos 107 (2a); 246 (2a); 402 (2a); Tillett, $8. & C
Tillett - 1S op (12); 45380 (7); Tillett, S. et al. 45118 (12).

Ule, E. 8721 (4); 8725 (11).

Valera, A. 563 (12); Valverde, L. & I. Pefia 1021 (2a); VEN Herb. No. 3879 (2a); Vogel, P. 177 (2a):
309 (2a); 1118 (12).

Warming, J. s.n. (1); 553 (2a); 560 (12); Weitzman, A. & N. Holbrook 183 (2a); Werff, H. van der &
P. Ortega 8082 (2a); Wertf, H. van der & R. Rivero 7838 (2a); 7859 (2a); 8009 (2a), Wer ff, H. va
der et al. 8779 (2a); Williams, L. 9938 (2a); 11125 (12); Williams, L. & A. Alston 314A (2a); or6
(2a); Wurdack, J. & J. Steyermark 1160 (11).

—

ADDITIONS TO THE GENUS ARDISIA
SUBGENUS GRAPHARDISIA (MYRSINACEAE)

JOHN J. PIPOLY III

Botanical Research Institute of Texas
509 Pecan Street, Fort Worth, TX 76102- q008. OSA:
Jpipoly @brit.org; clusia@latinmail.com

JON M. RICKETSON

Missouri Botanical Garden
P.O. Box 299, St. Louis, MO 63166-0299, U.S.A.

jon. vicketson @mobot.org

ABSTRACT

Revision of the group of species formerly placed in Zvnilia Lundell revealed that the
group is synonymous with Ardisia cae: Graphardisia Mez. This paper incorporates
those taxa in the subgenus, ee further revision of Graphardisia, including an emended
description of the subgenus, a key to the species and subspecies, Sail with updated de-
scriptions of the two added taxa a new illustrations for each. Two additional taxa ar
recognized, including one new combination: Ardisia verapazensis Rae cucullata Lund a
Pipoly & Ricketson. Six binomials are relegated to synonymy within A. verapazensis Donn
Sm. Ardisia hyalina Lundell {Zunilia en (Lundell) Lundell] is excluded from subgenus
Graphardisia, and placed in subgenus Ardisia

RESUMEN
Una revisi6n del grupo de especies anteriormente clasificadas dentro del género Zunilia
Lundell, revel6 que Zunilia es sinénimo del género Ardisia subgénero Graphardisia Mez.
Se incorporan dichos taxa dentro del subgénero ile donee y se ee una cesceipcion

actualizada, una clave para separar todos los tax
y nuevas ilustraciones para ellos. Se reconocen aes taxa adicionales, miley aaa la nueva
combinacién: Ardisia verapazensis subsp. cucullata (L ae Pipoly & Ricketson. Se relegan
seis binomios a la sinonimia bajo A. verapazensis Donn. Sm.. Se excluye Ardisia hyalina Lundell
[Zunilia hyalina (Lundell) Lundell} del subgénero Guia y se la ubica dentro del
subgénero Ardisia.

INTRODUCTION
In preparing our treatment of the Myrsinaceae for Flora Mesoamericana,
we critically reexamined taxa segregated by Lundell from Ardisia, includ-
ing Gentlea (Ricketson & Pipoly 1997), Graphardisia (Pipoly & Ricketson
1998a), Chontalesia and species related to it, now known to belong to the
genus Hymenandva A. DC. ex Spach. (Pipoly & Ricketson 1999). Since our
study of subgenus Graphardisia (Pipoly & Ricketson 1998a), it has become

Stipa 18(4): 11 1160. 1999

1146 Stipa 18(4)

evident that the taxa comprising the segregate genus Zvnilia (Lundell 1981)
are best placed within Ardisia subgenus Graphardisia.

In Lundell’s description of the genus Zuui/ia (Lundell 1981), he sepa-
rated it from the genus Ardisia based on the following key:

. Panicles of Howers in corymbs, the mature inflorescences heteromorphic with
strongly accrescent elongated pedicels arranged either in whorls or opposite
or alternate below the apical corymbs Zunilia

. Panicles of flowers consisting of simple racemes or spikes, the inflorescences

with flowers uniformly distributed, either pedicellate or sessile ..........00 ee Ardisia

However, we noted that Lundell failed to compare or distinguish Zilia
from his genus Graphardisia. As we studied the group of species assigned
by Lundell (1981) to Zunilia, we noted their raised or sessile glandular-
papillace filaments, the large accrescent, densely and prominently black punctate

and punctate-lineate sepals that clasped the developing fruit, the black punctations

and punctate-lineations of all abaxial leaf surfaces, the lanceolate, concolorous,
apiculate anthers with subapically poricidal dehiscence, and the style 2—3
times longer than the ovary, all features we used (Pipoly & Ricketson 1998a)
to define subgenus Graphardisia. Therefore, we find it necessary to relegate
the genus Zvnilia to synonymy under Ardisia subgenus Graphardisia, specifically
as an emended concept of Ardis1a verapazensis Donn. Sm.

NOTES ON KEYS, TAXONOMIC CONCEPTS, TERMINOLOGY

The key is artificial and designed to expedite identification of herbarium
specimens. An attempt has been made to emphasize vegetative characters
to increase the key’s usefulness with sterile material. The respective posi-
tions of taxa in the key and correlations with sence relationships are
coincidental. Quantitative and qualitative data presented in keys and de-
scriptions for Horal parts and bracts were taken from organs rehydrated from
herbarium specimens by boiling in water. Measurements from these range
from 10% to 15% greater than those measurements taken directly from
dried material. Data regarding stem diameters, inflorescence rachises, pedicels,
leaf and fruit shape and size were taken from dried herbarium specimens.

Our concept of subspecies follows that of Pipoly (1987) who defined subspecies
as: “groups of populations within a single lineage of ancestor-descendant

a

populations that show variation by unique combinations of plesiomorphies,
or homoplasic apomorphies, correlated with biogeography and/or ecology.
This rank is primarily used to convey information regarding variation 1n
the life histories of these populations and character state differences hypothesized
to be the result of this variation. The subspecific rank in no way attempts
to predict speciation events.”

Morphological terms in this treatment follow Lindley (1848) and Pipoly
(1987, 1992) for the inflorescence, rachis pedicels and floral parts. Descrip-

Pirpoty AND RickeTson, Additions to Ardisia subgenus Graphardisia LIA?

tion of leaf morphology follows Hickey (1984), trichome description fol-
lows Theobald et al. (1984) and basic cell and tissue terminology follow
Metcalfe (1984),

TAXONOMIC TREATMENT
Ardisia Subgenns Graphardisia Mez in Engl., PAlanzenr. IV. 236 (Heft 9):78.
1902; Lundell, Wrightia 3:192-198. 1966. Graphardisia (Mez) Lundell,
een 48:139. 1981; Lundell, Phytologia 59:429-433. 1986. Type: Ardisia opegrapha
Oerst.(LecToTYPE, by Lundell, Phytologia 48:139. 1981).
Zunilia Lundell, Phytologia 49:353. 1981, Syn. Nov. Type: Ardisia sexpartita Lundell,
Wrightia 3:29. 1962. Zunilia sexpartita (Lundell) Lundell, Phytologia 49:354. 1981.

Subshrubs to trees. Branchlets mostly terete, glabrous or rarely, glandular-
granulose. Leaves petiolate; blades membranaceous to subcoriaceous, densely
and conspicuously or inconspicuously black (rarely pellucid-) punctate and

punctate-lineate, the margins entire to crenulate, rarely sharply and irregularly
dentate. Inflorescence terminal, uni- to tripinnately paniculate, pyramidal to
obpyramidal, rarely globose, the ultimate branches corymbose, at times in
high anthotactic spirals and thus appearing umbellate, the rachis often densely
and prominently black punctate and punctate-lineate; inflorescence and floral
bracts foliaceous, mostly persistent, resembling the vegetative leaves but
acropetally reduced in size. F/owers with perianth white, pink, lavender or
purple, densely and prominently black punctate and punctate-lineate; ca-
lyx with sepals free or nearly free, large, accrescent and clasping fruits at
maturity; corolla rotate, the lobes imbricate in bud, basally short-connate
and sparsely to densely yellow glandular-granulose or papillose within; stamens
inserted at corolla tube base, the filaments basally connate to form an in-
conspicuous tube, the tube free from the corolla tube, the apically free por-
tions of the filaments glabrous or glandular-granulose or glandular-papil-
late, less than half the length of the anthers, the anthers ovate-lanceolate,
linear or lanceolate, prominently apiculate, dehiscent by subapical pores;
ovary globose to depressed-globose, the style slender, 2—3 times longer than
the ovary, the placenta apiculate, the ovules pluriseriate, biseriate, or ap-
parently uniseriate (few in number and ina uy high anthotactic spiral).
Fruit globose or oblongoid, densely conspicuously punctate and punctate-
lineate, usually basally surrounded by persistent, clasping sepals.

Bolivia and adjacent Brazil.

Ecology. enus occur in diverse vegetation types, including
wet and pluvial lowland, premontane, montane, and cloud forests.

The subgenus is defined by: 1) glabrous branchlets and inflorescence ra-
chises; 2) dense and prominently raised or conspicuous black, or rarely, reddish-
black, punctations or punctate-lineations on all leaf and/or floral parts; 3)

1148 Sipa 18(4)

sepals often accrescent and usually clasping the developing fruit; 5) linear-
lanceolate, concolorous, apiculate anthers with subapically poricidal dehis-
cence; and 6) style 2—3 times longer than the ovary. Species of the subgenus
are often used for home decoration and for use in Christian churches for
religious holidays (Pipoly, pers. obs.).

KEY TO TAXA OF ARDISIA SUBGENUS GRAPHARDISIA

1. Stoloniferous subshrubs mostly less than 1 (—2) m tall; leaf blade margins
sharply and irregularly dentate; corolla tube and filaments yellow glandular-
granulose; Ecuador to Bolivia and adjacent Brazi A. weberbaueri

1. Shrubs to small trees (0.5—)2—6(— ‘Ni )m tall without seolons? leaf blade margins
entire, undulate or crenulate; corolla tube yellow glandular- ass anulose; filaments
gl ibrous or sessile to stalked endl papillate; Mexico to Colombia.

. Sepals membranaceous, oblong, 4.2—8 mm lois apically broadly rounded
A.

to obtuse, hyaline throughout, the margins entire opegrapha
_ Inflorescence obpyramidal; leaf blades oblanceolate or rarely obovate,
3.5-7.5(—8) cm wide, 3 or more times longer than wide.
4. Floral bracts caducous; stamens 5.2—6.5 mm long; filaments 2.5—3
mm long; sepals S—8 mm eee A. opegrapha aes opegrapha
. Floral bracts cas: stamens 3.8—5 mm ee filaments 1.5
mm long; sepals 4.2-5.2 mm ic phaitueas inane . opeg aed subsp. wagner!
. Inflorescence g leboses on blades el lipeie ro broadly elliptic G (7.5-
wide, 2—2.5 times longer than wide. ..... A. opegrapha ee paquitensis
2. Sepals chartaceous, ovate, |.5—3.2(—4.0) mm long, apically acute to rounded,

opaque except at margin, the margins subentire to erose.

obes ovate, elliptic or lanceolate; filaments glabrous; ovules
; Panama and Colombia A. bartletti

7

6. Bea 1.5-1.8 mm long; petal lobes 6—6.5 mm long; stamens 3.5

5S. es

4.8 mm long; style base tapering; call wet forests. 00... A. bartlettit
aubep: bartletti
6. Sepals 2—-2.5 mm long; petal lobes 7-8 mm long; stamens 4.5—5.7
mm long; style base stylopodic; strand vegetation oe beach fore

. bartlettit ae lilacina

5. Corolla lobes oblong; filaments sessile- to oe yapillate;
ovules 22—35; Oaxaca, Mexico to Honduras : verapazensis

D

. Corolla lobes 6.5—6.7 mm long; free fae of filaments 3.3—3.4

rut —9 mm in diam., fruiting

mm long; style 3-3.1 mm long;
style 7—8.6 mm long, usually only oi basal portion persistent
verapazensis subsp. as ela:

7. Corolla lobes 5.7—5.9 mm long; free portion of filaments 2.6—2

mm long; style 5.5—5.9 mm long; fruit S—G.1 mm in diam., ae
ing style 8.8—-9.4 mm long, entire style usually persistent ...... A. verapazensis
subsp. cucullata

Ardisia verapazensis Donn. Sm.

Shrub or small tree to 30 m tall, 33 em DBH. Branchlets slender to stout,
(3—)5—8 mm in diam., glabrous. Leaves with blades membranaceous to

Pipoty AND RickeTson, Additions to Ardisia subgenus Graphardisia 1149

chartaceous, elliptic to oblong or oblanceolate, 7.4-34.5 cm long, 3.2—9.7
cm wide, apically acute to acuminate, basally acute to acuminate, decur-
rent on the petiole, midrib impressed above, prominently raised below, the
secondary veins prominulous above, conspicuous to inconspicuous below,
at times prominently reticulate, mostly inconspicuously punctate and punctate-
lineate, glabrous, the margins entire, undulate to regularly or irregularly
crenulate; petioles canaliculate or marginate, 0.4-1.4 cm long, glabrous.
Inflorescence terminal, bipinnately paniculate, 4.8-16.2 cm long, 3.9-22.4
cm wide, the rachis glabrous, the branches terminating in corymbs; peduncle
0.9-4.2 cm long; secondary inflorescence bracts early caducous, membranaceous,
ovate or oblong, (5.4—)9.9-14.8 cm long, 3.7-4.8 cm wide, apically acute
or rounded, otherwise similar to the leaves; floral bracts very early cadu-
cous, not leaving detectable scars axillant to pedicel (aborted at primordial
stage?), or early caducous, membranaceous, ovate, (..8—0.9 mm long, 0.7—
0.8 mm wide, apically rounded, the midrib inconspicuous, the secondary
veins not visible, densely and prominently black punctate and punctate-

—_—

lineate, the margins hyaline, sparsely glandular-ciliolate; pedicels stout or
slender, 0.4-1.8 cm long, accrescent in fruit or not, glabrous. Flowers 5(—
6)-merous, membranaceous, chartaceous to coriaceous, 8.4—10.2 mm long;
calyx with us free, broadly ovate, 3.4-4.0 mm long, the lobes 2.7—3.2
mm long, 2.3—3.0 mm wide, apically acute to rounded, conspicuously and
ares punctate and punctate-lineate, sparsely scattered glandular-
glandulose within near the base, often sparsely lepidote medially outside,

the margins hyaline, erose to entire or subentire, ciliolate with multicellu-
lar hairs; corolla rotate, 7.7—9.2 mm long, the tube 1.9—2.5 mm long, the
lobes oblong, 5.7—6.7 mm long, 3.3—4.1 mm wide, apically rounded, con-
spicuously and prominently punctate and punctate-lineate, yellow glandular-
glandulose and or papillose at base between corolla lobe and tube junction
and above staminal tube, otherwise glabrous; margins entire; stamens 5.9—
7.7 mm long, the filaments 3.4—5.0 mm long, united basally into a stami-
nal tube 0.7-1.6 mm long, the apically free portions 2.6—3.4 mm long,
0.5—-0.7 mm diam., slender or stout, epunctate, sessile or stalked glandu-
lar-papillate, the anthers lanceolate, 3.0—3.2 mm long, 1.2—1.4 mm wide
at base, apiculate, basally sagittate, dehiscent by subapical pores; ovary glabrous,
the style 3.0-5.9 mm long, slender, inconspicuously punctate and punc-
tate-lineate, glabrous, the ovules 22—35, pluriseriate. Frit globose to de-
pressed-globose, S—9 mm in diam., style 7—9.4 mm long, persistent at least
basally, densely and conspicuously punctate, glabrous.

Within subgenus Graphardisia, Ardisia verapazensis is most closely related
to Ardisia opegrapha, because of its large, foliaceous inflorescence bracts and
glandular-papillate filaments.

1150 Sipa 184)

la. Ardisia verapazensis Donn. Sm. subsp. verapazensis, (Fig. 1, 2). Andisia
verapazensis Donn. Sm 7 PDE Gaz. 46:113. 1908. Zunilia verapazensis (Donn. Sm.
Lundell, Phytologia - 49:3 4. LOSl. Type: GUATEMALA. Attra VeERApaz: In monte
silvoso prope Coban, co m, Jan 1908 (fl), H. von Tiirckheim 12093 (HOLOTYPE:
US; isotypes: BR, F 2-sheets, G, GH, MO, NY, US 2-sheets).

Ardisia sexpartita Lundell, Wrightia 3:29. 1962, syN. Nov. Zunilia sexpartita (Lundell)
Lundell, Phytologia - 49:354. 1981. Type: GUATEMALA. QuEZALTENANGO: lower S-
facing cs of Volcdin Santa Maria, near San Juan Patzulin, 1,300-1,500 m, 6 Jan

ne 4 — ) ¥? aon 33608 (MoLoTyYPE: LL).

\y intlensis Lundell, Wrightia 3:98. 1964, syn. Nov. Type: GUATEMALA. EscuINtLa:
veut further localiy without elev., 1942 (fl), J. Lenacio Aguilar 1679 (HOLOTYPE: F).

Ardista alba Lundell, Wrightia 3:195. 1966, syn. Nov. Zunilia alba (Lundell) Lundell,

hycologia 49:353. 1981. Typr: MEXICO. Cuiapas: Municipio Jitotol, steep wooded
slope on bank of Rio Hondo, 4 mi N of Jitotol on road to Pueblo Nuevo Solistahuacan,
5,500 fe {1,676 m}, 12 Feb 1965 (A), D. Breedlove 8962 (HoLorypE: LL; isorypes: DS,

MICH).
sal gis Lundell, Phytologia 58:490. 1985, syn. Nov. Ardisia eciliata (Lundell)
undell, Phytologia 61 oe . 1986, (nomen invalidum). Ardisia eciliata (Lundell) Pipoly

& soar on. Sida 18:512. 1998. Type: MEXICO. Cutapas: Municipio Ocozocoautla
de Espinosa, steep ops i; Cerro del Ocote, 30 km NW of Ocozocoautla, montane

rain forest, 1,500 m, 14 Oct 1972 (A), D. Breedlove 28973 (uoLoryer: LL; YPES:
DS, MEXU, MICH, ve

Zunilia parpasis Lande ll, Phytologia 58:491. 1985, syn. Nov., non Ardisa 77 Brandeg.,
Univ. Calif. Publ. Bot. 6:189. 1915. Ardisia feniana ees l, Phyo 26k 64. 1986,
(nomen invalidum). Ardisia fentana Pipoly & Ricketson, Sida 18:512 1998. TYPE:

MEXICO. Cniapas: Mountains near Fenia, without elev., May ee (fr), C ». Purpus
100 (HOLOTYPE:

Ae

Shrub or small tree to 30 m tall, 33 cm DBH. nea stout, (3—)6—§
mm in diam. Leaves with blades 7.4—34.5 cm long, 3.2—9.7 cm wide, the
secondary veins inconspicuous below, the margins entire, ances to regularly
crenulate; petioles marginate, 0.6-1.3 cm long. Inflorescence 4.8-16.2 cm
long, 3.9—22.4 cm wide; peduncle 1.3—4.2 cm long; secondary inflorescence
bracts early caducous, membranaceous, ovate or oblong, (5.4—)9.9-14.8 cm
long, 3.7-4.8 cm wide, apically acute or rounded, otherwise similar to the
leaves; floral bracts very early caducous, not leaving detectable scars axillant
to pedicel (aborted at primordial stage?); pedicels stout, 4-18 cm long, accrescent
in fruit. F/owers chartaceous to coriaceous, 10—10.2 mm long; calyx 3.7—
4.0 mm long, the lobes 2.8—3.0 mm long, 2.6—3.0 mm wide, often lepi-
dote medially without; corolla 9.0—9.2 mm long, the tube 2.4—2.5 mm long,
the lobes 6.5—6.7 mm long, 3.8—4.1 mm wide; stamens 7.5—7.7 m long,
the flaments 4.8—5.0 mm long, united basally into a staminal tube 1.5—
1.6 mm long, the apically free portions 3.3—3.4 mm long, 0.5—0.6 mm diam.,
slender, scattered yellow stalked or rarely sessile glandular-papillate, the
anthers 3.0—3.2 mm long, 1.2—-1.4 mm wide at base; style 3.0—3.1 mm

jes

Pipoty AND RickKEeTsON, Additions to Ardisia subgenus Graphardisia 1151

Fic. |. Ardisia verapazensis subsp. verapazensis, showing foliaceous secondary inflorescence
bracts enclosing the immature inflorescence. Drawn from A. Méndez Ton 4488 (MO).

1152 Sipa 18(4)

Fic. 2. Ardisia verapazensis Donn. Sm. subsp. verapazensis. A. Reproductive shoot. B. Detail
of abaxial leaf surface. C. Detail of inflorescence. D. Detail of flower. E. Detail of stamens,
showing adaxial surface. F. Detail of stamens, showing abaxial surface. G. Detail of sta-
mens, showing laterial surface. H. Fruit. A-H drawn from H. von Tiirckheim 12093 (GH
isotype).

PipoLy AND RicKETSON, Additions to Ardisia subgenus Graphardisia Lie

long; ovules 22-26. Fruit 6.5—9 mm in diam.; style 7-8.6 mm long, usu-
ally only basal portion of style persistent.

Distribution.—Ardisia verapazensis subsp. verapazensis is known from Oaxaca,
Mexico through central Chiapas to Guatemala (Alta Verapaz, Quiché,
Huehuetenango, Quezaltenango, San Marcos, Suchitepéquez) and with one
disjunct population in Copdn, Honduras. It grows from 5O00—3,000 m el-
evation.

Ecology and conservation status —Ardisia verapazensis subsp. verapazensis 1s
known from primary premontane and montane wet forests, and cloud for-
est margins. It is locally common, but not known to grow in large popula-
tions. Given its relatively wide ecological tolerance, this species is not con-
sidered threatened at this time.

Etymology. —The epithet ‘verapazensis’ refers to the type locality, near Coban,
Department of Verapaz (now Alta Verapaz) in northern Guatemala.

Local names.—’ Shyash tililja” (Tzeltal), (A. Méndez T’ 4404), “niwektililjaz”
(Tzeltal), A. Méndez T. 4488); “b’shyashtililjas,” (A. Shilom T. 7451); “huesito
blanco,” (J. Steyermark 3363 1); “palo cruz,” (J. Steyermark 37367); “camaco,”
(T. Hawkins & D. Mejia 241).

Specimens examined. MEXICO. Chiapas: ee Berriozabal, 13 km N of Berriozabal
near Pozo Turipache and Finca El Suspiro, 900 m, 71 (fr), D. Breedlove 20251 (DS,
LL, MICH, MO, NY), 1,000 m, 2 Nov 1971 (fr), D. = ae G " Smith 21680 (DS, LL,
MICH, MO), 900 m, 26 Dec 1972 (fr), D. Breedlove & R. Thorne 30777 (DS, LL, MICH,
M , TEX), 900 m, 15 May 1973 (fr), D. Breedlove 35294 (DS, LL, MICH, MO, NY),
1,000 m, 10 Oct 1980 (fr), D. Breedlove & J. Strother 46019 (CAS, LL, MO), 1,020 m, 28
Sep 1988 (fl, fr), D. Breedlove 70218 (CAS); SE side of Volcan Tacana above Talquian, 2,200
m, 16 Jan 1973 (fr), D. Breedlove & A. Smith 31707 (DS, LL, MICH, MO) Cerca de El Suspiro,
al NNW de Berriozabal, 900 m, 6 Sep 1953 (fl), EF Miranda 7887 (DS, US); Municipio
Bochil, along the river E of Bochil, 5,200 ft [1,585 m], 27 Jun 1967 (fr), A. Shilom T: 2574
(DS, F, LL, MICH); Municipio Cint ae a, ridge SE of Cerro Baul on the border of the state
of Oaxaca, 16 km NW of Rizo de Oro along a logging road to Colonia Figaroa, 1,600 m,
27 Apr 1972 (fr), D. Breedlove 24916 (DS), On Oaxaca-Chiapas border near La Cienega de
Leon, 1,080—1,230 m, | Dec 1980 (fr), D. Breedlove G F. Almeda 48217 (CAS, GH, LL,
NY); Municipio Pueblo Nuevo Solistahuacdan, near Clinica Terba Buena, 3 km NW of Pueblo
Nuevo Solistahuacan, 1,700 m, 14 Dec 1971 (fr), D. Breedlove 23201 (DS, MICH, MO),
2.5 mi N of Pueblo Nuevo Solistahuacan, 5,300 ft {1,615 m}, 19 Jun 1970 (fr), E. Lathrop
GR. Thorne 7233 (DS), near Clinica Terba Buena, 2 km NW of Pueblo Nuevo Solistahuacdn,
5,400 ft {1,646 m}, 23-24 Jan 1965 (fr), P. Raven & D. Breedlove 19918 (DS, F, MICH,

US), 3 km NW of Pueblo Nuevo Solistahuacan, on the slopes below Highway 195 in the
vicinity of Clinica Yerba B , 17° 30" N, 92° 40" W, 5,400 fr (1,646 m), 7 Oct 1971
(fr), R. Thorne & E. Lathrop “46084 (MICH, NY), 17° 30" N, 92° 40" W, 5,800 ft [1,646
ml, 20 Sep 1970 (fl), H. Zuill 398 (DS); 9 Oct 1970 (A), H. Zuill 650 (DS), ennnean
Rayon, in the Selva Negra, 10 km above Rayén, Mezcalapa, along road to Jitotol,

m, 13 Jul 1972 (fr), D. Pili 26072 (DS, LL, MICH, MO, NY), 10 Jan ne) a bud,
D. Breedlove & B. Keller 49316 oes LL, MO, NY), 9 mi NW of Pueblo N

along the road between Rincon Chamula and Rayé6n, slope near Puerto del Viento, 17° 30"

N, 93° 40" W, 5, 0 a a ar Sep 1971 (fl), R. Thorne & E. Eanes (DS); Municipio

—

No)

1154 Sipa 18(4)

San Cristobal de Las Casas, Santa Cruz en San Filipe, without elev., 15 Nov 1986 (fr), A.
Méndez T. & M. C. Martinez. de Lopez 9844 (CAS, MO, NY, © EX): Municipio Tenejapa,
near Paraje Yashanal, 2,460 m, 28 Jan 1981 (fr), D. Breedlove 49645 (CAS), 2,300 m 22
me 1981 (fr), D. Breedlove 51046 (CAS); 1,980 m, 13 Jul 1981 (fr), D. Breedlove 51463
LL); Municipio Tenejapa, Ojo del Rio Yashanal, 1,700 m, 10 Jul 1982 (fr), A. Méndez
a Ps MO), 20 Aug 1982 (fl-bud), A. Méndez T. 4488 (MEXU, MO); Municipio
Tila, Colonia Kokijaz, 1,000 m, 20 Mar 1983 (fr), A. Méndez T. 5700 (MEXU, MO), 5
May 1983 (fr), 5965 (MEXU, MO, TEX), Finca eerie without elev., 20 Mar 1984 (fr),
- Shilom T. 7451 (LL, MEXU, MO); Mt. Tacana, 2,000—4,038 m, Aug | ene E. Matuda
392 (FL GH, LL, MICH, MO, NY, UC, US); Barr. Alpujarrez, without elev., 4]
i FE. Miranda 1746 (MEXU, TEX); Mes. near Fenix, without elev., pee ee 930 (fr),
C. Purpus 10100 (GH, NY);. Oaxaca: Distrito Ixtlan: Municipio Comaltepec, SW slope of
Cerro muna be above eee in, near Federal Elect cEreity Commission Camp,
near Highway 17 7° 29' 15" N, 96° 24! 05" W, 1,750—1,780 m, 3 Dec 1993 (ster.), B.
Boyle et al. 2643 min 15 mi of San Gabriel along road from Puerto Escondido to Oaxaca,
6,000 fr (1,829 m), 9 May 1965 (fr), D. Breedlove 9880 (LL), La Esperanza, 17° 37'N, oo
21'W, ca. 1,600 m, 9 Jan 1988 (fr), R. Lopez L. & G. Martin 193 (MQ), 17° 37'N, 96° 21
W, 1,600 m, 27 Jul 1989 (fl), R. Lopez L. & G. Martin 491 (MO), Vista Hermosa, trail to
camp, 48.8 km SW of Valle Nacional, 17° 39'N, 96° 19' W, 1,460 m, 23 Jan 1988 (fr), R.
Torres C. & L. Cortes 11623 (BRIT, MEXU); Distrito Miahuatlan: Municipio San Jer6nimo
nae an, 18 km NE of Piedra Larga, road to San Jer6nimo Coatlan, 16° 09'N, 97° O1' W,
1,950 m, 16 Jan 1988 (fr), A. a Vv. on - MEXU, MO), 11.5 km SW of San ee
Coatlan, trail to Piedra Larga, 16° 20' N, 96° 57' W, 2,050 m, 17 May 1988 (fr), A. Cam-
pos V. 1819 (EK MEXU, ae spel de a. 13.5 km SW « song aah an An tie
trail to Piedra oS 16° 12'N, 96° 57' W, 1,950 m, 17 May 1988 (fr), A. Campos V. 1834
(F, MEXU, MO), 17.9 - SW of San Jerénimo Coatlan, trail to Piedra | 16° 12'N,
96° 58' W, 1,890 m, 17 May 1988 (fr), A. Campos V. 1853 (F, MEXU, MO), 19.2 km SW
of San Jer6nimo Coatlan, trail to Piedra Larga, 16° 12! N, 96° 58! a 1,900 m, 13 Aug
1988 (fl, bud), A. Campos V. & L. Cortés 2242 (F, MEXU, MO); 6 km NE of logging camp
Cerro Sol, trail to Progreso, 16° 11' N, 97° 00! W, 1,400 m, 6 Oct 1988 (fl), A. Campos V.
2585 (F, MEXU, MO); 20 km W of San Jerénimo Coatlan, 1,680 m, 26 Oct 1982 (A, fr).
E. Martinez et al. 2501 (BRIT, MEXU); page he Nacional, Se ee 15 km
NNW of Valle Nacional, 17° 93' N, 96° 'W, —1,200 m, 15 Oct 1992 (fr), J.
Meave astillo et al. 1491 (MEXU, MO), Ca etal t Lucfa, 1,300 m, 9 | ake) (A),
B.Reko 441 (US), 20.5 km SW of San Jeronimo Coatlan, trail to Pied Larga, 16° 12
96° 58! ae 2,000 m, 15 Mar 1989 (fr), G. Toriz A. & A. Campos V. 818 (FE, MEXU, MO).
Cerro Batl, 23 km NE of Rizo de Oro, road to Colonia Rodolfo Figueroa, without elev., 28
Mar 1984 (fr), R. Torres C. GC. Martinez ¢ iat (MEXU, MO); 9.6 km SE we erro de Vidrio,
Oaxaca-Puerto Escondido Highway, 1,850 m, | Aug 1984 (fr), R. Torres C. & C. Martinez
5824 (MEXU, MO); Distrito Mixe Mian un icipio Totontepec, 10.5 km N of Toomer road
to Choapan, 17° 17! 00" N, 95° 59! 00" W, 1,760 m, 27 Oct 1988 (Al, fr), R. Torr L.
Cortes 10388 (BRIT, MEXU), ween San Miguel Chimalapa, Cerro Salomo1 n
in one at line NNW of Cerro Guayabitos, ca. 43 km in straight line N of San Pedro fears
16° 45'N, 94° 11' 30" W, 1,850 m, 23 ne 1985 (fl, fr), 7 Wendt et al. 5149 (BRIT, LL,
MEXU); Distrito Sola de Vega, Sean Santa Cruz Zenzontepec, El Carrizal, 16° Cn
75" N, 97° 26' 06" W, 1,040 m, 14 Apr 1993 (ster.) J. Weiss 118 (TEX). GUATEM
Alta Verapaz-Quiché: Chama to an 3,000 ft [914 m}, 15 Oct 1920 (A), H. aan
S65 (EF, LL-fragment, US). Huehuetenango: Cerro Huitz, between Mimanhuitz and Yulhuitz,
Sierra de los Cuchumatanes, 1,500—2,600 m, 14 Jul 1942 (fr), J. Steyermark 48602 (F, US).

PrpoLty AND Ricketson, Additions to Ardisia subgenus Graphardisia BSS)

Quezaltenango: Volcan Zunil, 5,500 ft {1,676 m}, 3 Aug 1934 (fl), A. Skutch 931 (F, GH),
Volcdn Santa Maria, between Santa Marfa de Jestis and Calahuaché, along great barranco
between Finca Pirineos and San Juan Patzulin, slopes at San Juan, 1,300-1,500 m, 6 Jan
1940 (ster.), J. Steyermark fone (F); Between Quebrada Chicharro and Montafia Chicharro
on SE-facing slopes of Volcan Santa Marfa, 1,300—1,400 m, 18 Jan 1940 (fr), J. Se
34363 (F). San Marcos: ae Armenia, Reed de Cuesta, ca, 5,000 ft [1,524 ml}, 6-7 Jul
1977 (fr), J. Dwyer 14458 (LL, MO); Rio Vega, near San Rafael and Guatemala-Mexico
boundary, Volcan Tacand, 2,500—3,000 m, 20 Feb 1940 (fr), J. Steyermark 36261 (E); Above

Finca El Porvenir, up Loma Bandera Shac, lower S-fi olcan Tajumulco, 1,300—
1,500 m, 9 Mai 1940 (fr), J. Steyermarke 37367 (F, LL Near Aldea Fraternidad, between
San Rafael Pie de la Cuesta and Palo Gordo, W-faci of the Sierra Madre Moun-

tains, 1,800—2,400 m, 10-18 Dec 1963 (fl), L. vie et | 25769 (BM, F, UC), 26085
(F, NY), 26278 (E&, GH, US); Outer ee of Tajumulco Volcano, Sierra Madre Mountains
about 8-10 km W of San Marcos, ca. 2,300 m, 31 Dec 1964—1 Jan ne (fl), L. Williams
et al. 26799 (FE, NY, US), (fr), L. Williams et al, 27188 (F). Suchitepéquez: Slopes of Volcan
Zunil, eae, of Finca Las Nubes, along Quebrada Chita, E of Pueblo Nuevo, 500-800
m, 2 Feb 1940 (fl), J. Steyermark 35431 (F), Volcén Santa Clara, between Finca El Naranjo
and upper slopes, 1,250—2,650 m, 23 May 1942 (fr), J. pes 46650 (F, US). Without
department: Las Nubes, without elev., Nov 1877 (fH, fr), K. Ber noullt & A. Cario 1975
(GOET). HONDURAS. Copan: S slope of Cerro Azul, 12 km NW of Florida, Cerro Azul
National Park, 15° 6' N, 88° 55' W, 1,500 m, 11 Feb 1992 (fr), 7) Hawkins G D. Mejia
241 (EAP, HEH, MO, TEFH)

Ardisia verapazensis subsp. verapazensis exhibits great quantitative varia-
tion among relative size of its organs within individuals and among popu-
lations. Even though its distribution is somewhat restricted, this had led
to taxonomic overdescription, much like what Pipoly and Ricketson (1998a)
noted for species such as Ardisia opegrapha Oetst.

The type of Ardisia sexpartita was collected from populations whose
inflorescences are smaller than average and whose flowers are more waxy
pink than the average. Ardisia escuintlensis’s type is a fragmentary collection
whose flowers (in bud) have thinner perianth parts, but whose organs oth-
erwise fall well within the size range of variation for the taxon. Ardisia alba
was described only because of its white flowers and inflorescences that dry
almost white, its relatively larger anthers and less conspicuous punctations.
The holotype of Zunilia eciliata, mostly in fruit, is notable for anthers slightly
smaller than the average for the species, and calyx lobe margins that are
entire and without glandular cilia. However, reexamination of the MO isotype
clearly shows small scattered cilia present on young calyx lobes in bud. Zunilia
purpusii, whose type is a fruiting collection, was separated by its larger ca-
lyx lobes with acutish apices. However, both of these features lie well within
the range of variation for the subspecies.

It is notable that Lundell’s combinations of Ardisza eciliata (Lundell) Lundell
and Ardisia feniana (Lundell) Lundell are both invalid names. We subse-
quently validated these names with our combinations Ardisia eciliata (Lundell)

ee

1156 Sipa 184

Pipoly & Ricketson and Ardisia feniana (Lundell) Pipoly & Ricketson (Pipoly
& Ricketson 1998b). However, our current studies show that these taxa are
clearly synonyms of Ardisia verapazensis subsp. verapazensis.

a

Ib. Ardisia verapazensis Donn. Sm. subsp. cucullata (Lundell) Pipoly &
Seascale. emp cf vole nov. (Fig. 3). Ardisia cucullata Lundell, Wrightia
3:26. 1962 1 idell) Lundell! Phy tologia 49:354, 1981. Type: MEXICO.
c HIAPAS: a lesca, near or See 2,000 m, 7 Mar 1945 (fl), E. Matuda 5201 (HOLO-
2 LL; tsorypes: FE J).

oa
~—

=

Zunilia mirandae Lundell, ee ogia 38:491. 1985, sYN. NOv., non Ardisia mirandae Merr.,
Philipp. J. Sct. 12:154. 1917. Ardisia neomirandae Lundell, Pt He ia GL 1986,
eae invades: ie anna Pipoly & Ricketson, Sida 18:5 14 1998. TYPE:
MENICO. Cutapas: Finca Prusia, along road from Mapastepec to near Ti ie jie
without elev., 24 Feb 1951 (fH), EK Miranda 7004 (HoLoryPE: MEXU),.

Shrub or small trees to 12 m tall. Branchlets slender, 3.5—5(—7) mm in diam.
Leaves with blades 9.2—20.6 cm long, 3.5—8.2 cm wide, the secondary veins
conspicuous below, prominently reticulate, che margins entire to undulate,
to irregularly crenulate; petioles canaliculate, slender, 0.4—-1.4 cm long. Inflorescence
9.S-11.5 cm long, 7-15 cm wide; peduncles 0.9-2.5 cm long; secondary
inflorescence bracts early caducous, unknown; floral bracts early caducous,
membranaceous, ovate, minute, 0.8—0.9 mm long, 0.7—0.8 mm wide, apically
rounded, the midrib inconspicuous, the secondary veins not visible, densely

and prominently black punctate and punctate-lineate, glabrous, the mar-
gins entire, hyaline, sparsely glandular-ciliolate; pedicels slender, 0.8—1.7
cm long, not accrescent. Flowers esas 8.4-8.6 mm long; calyx

.4—3.9 mm long, the lobes 2.7—3.2 mm long, ee 5 mm wide, glabrous
hoa corolla 7.7—7.9 mm long, the tube 1.9—2.1 mm long, the lobes
5.7-5.9 mm long, 3.3—3.5 mm wide; stamens 5.9—6.0 mm long, the fila-
ments 3.4—3.6 mm ee united basally into a staminal tube 0.7—0.9 mm
long, the free portion 2.6—2.8 mm long, 0.6—0.7 mm in diam., stalked or
rarely sessile glandular-papillate, the anthers, 3.0—3.1 mm lone: 1.3-1.4
mm wide at base; or 5.5-5.9 mm, ovules 10-16. Frait 5.0—6.1 mm in
diam., style 8.8—9.4 mm ie entire style usually persistent.

Distribanen —Ardisia verapazensis subsp. cucullata is endemic to the south-
western region of Chiapas, Mexico, around the Sierra Madre de Chiapas and
the Area Natural de Reserva E] Triunfo. It grows from 700—2,540 m elevation.

Ecology and conservation status.—This subspecies grows in wet montane
and cloud forests. Alchough its geographic range is very restricted, it ap-
pears to be common, especially in the Reserva de El Triunfo, where it is
presumably protected.

Etymology. —The epithet ‘cucullata’ refers to what Lundell (1981) inter-
preted as hooded inner petals. The misinterpretation was based on Lundell’s
belief that the corolla was fused at the base into a tube, with two larger

Prpoty AND RicketTson, Additions to Ardisia subgenus Graphardisia

1157

ES

Ae

QL N
Ras SS
a Saw
CWA

N
a PS
a enn OX SO
Te
Selle ea
a aii

ED “NG Wan Ae

Fic.

3. Ardisia verapazensis Donn. Sm. subsp. ce,

wllata (Lundell) Pipoly & Ricketson. A.
Reproductive branchlet. B. Detail of abaxial leaf surface. C. Detail of inflorescence. D. Detail
of stamens, showing adaxial surface. E. Detail of stamens, showing abaxial surface. F. De-
tail of stamens, showing lateral surface. G. Fruit. A

—F drawn from E. Matuda 5201 (F isotype).
G drawn from R. Hapshire et al. 500 (MEXU).

L158 Stipa 18(4)

outer lobes and three inner lobes that were “cucullate” or hooded at the
apex. Reexé ation of the type revealed that the corolla is simply imbri-
cate and any hooded appearence comes from drying effects.

Specimens examined. MEXICO. weer es Angel Albino Corzo, along Rio
C ee near Finca C eli a 2,400 ft [732 m}, 26 Mar 1968 (fr), A. Shilom T.
3867 (DS, F, LE 2-sheets, MICI Y). ee Reserva del Triunfo, Cafiada
del Pavon NW of Triunfo, on m, 10 May 1982 (fr), J. Calzada et al. 8739 (EE) trail
NNW from El Triunfo e amp to ; alo Gordo Camp, | km from El Triunfo Camp, 15° 39!
N, 92° 50! W, 2,000 m ) Feb 990 (fr), R. Hampshire et al. SOO (BM, MEXU), 1-3 km
from El Triunfo Camp, . 39! : 92° 50' W, 2,000 m, 21 Feb 1990 (A, fr), R. Hampshire
etal, 519 (BM 2-sheets, MEXU, NY); trail WSW from Palo Gordo towards Finca Catarina,
15° 40'N, 92° SI! W, 2,000 m, 25 Feb 1990 (fr), R. He saa et al. 699 (BM moe
MEXU : Municipio Jaltenango-Mapastapec, Reserva E] Triunfo, near HQ, 15° 39'N, 92°
48' W, 1,900 m, May 1989 (fr), M. Heath G A. Long 31 a CHIP), Cafiada a near
Camp Headquarters, 15° 39' N, 92° 48' W, 1,400 m, May 1989 (fr), M. Hea th & A. Long
40 (CAS, CHIP); Municipio Mapastepec, bee de Reserva E] Triunfo, Cabal Eiond.
7 km S$ of camp, 1,500 m, | Mar 1990 (fr), / Shee et al. 1080 (BM 2-sheets, MO),
Canada Honda, 7 km $ of camp, 1,500 m, O01 Mar (fr), A. Reyes G. et al. 1080 (BM 2-
sheets, MO); Municipio Motozintla de Mendoza, steep canyon, SW side of Cerro Mozotal,
11 km NW of che junction of the road co Morozintla along road to El Porvenir and Siltepec,
2,100 m, 22 Nov 1976 (fr), D. Breedlove — (DS); 23 oe 981 (A), D. Breedlove & B.
Barbee — Wats) ere Siltepec, on ridge above Siltepec along road to Huixtla,
400 feb 1982 (fr), 1 ye . Ei Aone: 58249 (CAS, LL, MO, NY);
ace ese os 29 Dec on ((), LE, Matuda 393 (LL, US); Boquerén, Motovibtla,
2,450—2,540 m, 5 May 1945 (fr), E. Matuda 5416 (F, LL 3-sheets); Mt. Ovando, Escuintla,
without elev., 14 Nov 1945 (fl), E. Matvda 16228 (MO, US).

ion

Qu

Zunilia mirandae Lundell is known only from the holotype. It is notable
only for its short petioles, smaller inflorescence and smaller sepals, but is
otherwise indistinguishable the type of the subspecies.

Subspecies cucu//ata is distinguished from subspecies verapazensis by its
shorter corolla lobes and free portion of the filaments, the style much longer
in flower and fruic, and smaller fruit. It appears to be geographically iso-
lated, occurring only in the southwestern region of Chiapas.

EXCLUDED NAME
Ardisia hyalina Lundell, Wrigl qtia 3:99. 1964. Zunilia hyalina (Lundell) Lundell,
Phytologia 49:354. 1981. Type: MEXIC O. SAN Luis Poros: vicinity of Xilitla, Cerro
Miramar, 4,400 ft (1,531 m), 15 Jul 1947 (fl), Ru. Newman 19 (HOLOTYPE: US
The ovate anthers with subapical pores, opening into slits, and panicu-
late inflorescences bearing racemose branchlets, all indicate that this spe-
cies belongs to Ardisia subgenus Ardisia, and not subgenus Graphardisia.

ACKNOWLEDGMENTS

We thank the Missouri: Botanical Garden and the Flora Mesoamericana
Project, for funding that allowed J. Ricketson (MO) to travel to Fort Worth,

PipoLy AND RICKETSON, Additions to Ardisia subgenus Graphardisia L159

where he was joined by J. Pipoly (BRIT) on visits to the C.L. Lundell Her-
barium (LL), housed at the University of Texas at Austin. Without access
to that critical collection, assembled by C.L. Lundell over a period of nearly
60 years, the present study would not have been possible. We also thank
the curators of BM, BR, BRIT, CAS, CHIP, DS, EAP, F G, GH, GOET,
HEH, MEXU, MICH, MO, NY, TEFH, UC, and US for loans of speci-
mens. We are grateful to the staff of TEX, especially Billie L. Turner, Tom
Wendt, Carol Todzia, Beryl Simpson, and José Panero, for their coopera-
tion and hospitality. We are also grateful to the Project for subsidizing vis-
its of John Pipoly to MO to work with Jon Ricketson. We owe gratitude to
those who have been so instrumental in assisting us in our work, including
Gerrit and Jeany Davidse, Linda Oestry, Mary Bard, and Catherine Mayo,
(MO), Barney Lipscomb, and Jim Rivers (BRIT). Illustrations were pre-
pared by the junior author. Reviews of the manuscript by Gerrit Davidse
(MO) and Richard Rabeler (MICH), and meticulous copy editing by Barney
Lipscomb, greatly improved the presentation of the paper.

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RICKETSON, J. and J. Prpoty. 1997. A synopsis of the genus Gentlea (seoramucea’s and a key
o the genera Myrsinacee in Orne. Sida 17:6
Tio, W.L., RAHULIK, and R.C. Roiins. 1984. Trichome aeesnee and classification.

At CARS eae and L. Chalk, eds. ce of the dictyledons. Vol. 1. Systematic
anatomy of leaf and stem, with a brief history of the subject. Clarendon Press. Oxford,

U.K. Pp. 40-53.

1160 Sipa 18(4)
NUMERICAL LIST OF TAXA
la. Ardisia verapazensis Donn Sm. subsp. verapazensis
Lb. Ardisia verapazensis Donn. Sm. subsp. cucu/lata (Lundell) Pipoly & Ricketson
LIST OF EXSICCATAE

The figures in parentheses refer to the numbers from the numerical list
of taxa. Collection numbers in boldface type indicate type specimens.
Bernoulli, K. & A. Cario 1975 (la); Boyle, B. et al. 2643 (la); Breedlove, D., 8962 (la),
9880 (La); 20251 (1a); 23201 (1a); 24916 (1a); 26072 (1a); 28973 (1a); 35294 (1a); 41758
(1b); 49645 (1a); 51046 (La); 51463 (la); 70218 (la); Breedlove, D. & FE. Almeda 48217
(la); 58249 (1b); Breedlove, D. & B. Bartholomew 35766 (1b); Breedlove, D. & B. Keller
49316 (la); Breedlove, D. & A. Smith 21680 (1a); 31707 (la); Breedlove, D. & J. Strother
46019 (la); Breedlove, D. & R. Thorne 30777 (la)

Calzada, J. et al.8739 (1b); Campos V., A. eae 1819 (la); 1834 (la); 1853 Cla); 2585
(la); Campos V., A. & L. Cortés 2242

Dwyer, J., 14458 (la).

Hampshire, R. ec al. SOO (1b); 519 (1b); i (1b); L680 (1b); Hawkins, T. & D. Mejia 241
(la); Heath, M. & A. Long 31 (1b); 40 (

Ignacio Aguilar, J., 1679 (La).

Johnson, H., 865 (1a).

Lathrop, E. & R. Thorne 7233 (1a); Lopez L., R. & G. Martin 193 (la); 491 (la).
Martinez, E. et al. 2501 (1a); Matuda, E., 393 (1b); 2392 (la); 5201 (1b); 5416 (1b); 16228
(1b); 2392 (1a); Meave del Castillo, J. et al. 1491 (la); Méndez T., A., 4404 (la); 4488
(1a); 5700 (La); 5965 (la); Méndez T., A. & M. C. Martinez. de Lépez 9844 (1a); Miranda,
F., 1746 (la); 7004 (1b); 7887 (La).

Purpus, C., 100 (1a); 10100 (La).

Raven, P. & D. Breedlove 19918 (La); Reko, B., 441 (la); Reyes G., A. et al. 1680 (1b).
Shilom T., A., 2574 (la); 3867 sap 451 (la); Skutch, A., 931 (1a); Steyermark, J., 33608
(la); 33631 (la); 34363 (la); 35 (la); 36261 (la); nae (la); 46650 (La); 48602 (La).
Thorne, R. & E. Lathrop 41689 (la); 46084 (la); Toriz A., G. y A. Campos V. 818 (1a);
Torres C., R. & C. Martinez 4869 (1a): 5824 (la); Torres C. R. & L. Cortés 10388 (La);
11623 (la); Tiirckheim, H. von, 112093 (la).

Wendt, T. et al. 3149 (la); Williams, L. et al. 25769 (la); 26085 (1a); 26278 (la); 26799
(La)

Zuill, H., 398 (1a); 650 (La).

NOTES ON SECTIONAL DELIMITATIONS
IN ERIGERON (ASTERACEAE: ASTEREAE)

GUY L. NESOM
Biota of North America Program—North Carolina Botanical Garden
Coker Hall CB 3280
University of North Carolina
Chapel Hill, NC 27599, U.S.A.
RICHARD D. NOYES
Plant Genome Mapping Laboratory
Riverbend Research Center, Room 162
University of Georgia
Athens, GA 30602, U.S.A.

ABSTRACT

om,

Erigeron sect. Linearifolii is narrowed to a monotypic taxon, comprising only Erigeron
hyssopifolius of northern North America. Er/geron sect. Pycnophyllum is restricted to the 14
species of the E. folzosus group, which are primarily Californian in Shean The Mexi-
can species E. exi/is is transferred from sect. Crncinnactis to sect. Lamprocaules. Erigeron rhizomatus
(New Mexico) and E. /epidopodus (Chihuahua and Durango, Mexico) are eonioli tel as Evi igeron
sect. Geronpternix Nesom & Noyes, sect. nov. The two species are distinct from other eri-
gerons in their scaled-leaved, rhizomiform caudex branches, large, solitary heads, and long,
narrowly oblong cypselas. Sect. Geroupternix is the most primitive phylogenetic element of

Erigeron and of subtribe Conyzinae.
RESUMEN

Erigeron sect. Linearifolii se reduce a un taxon monotipico, que comprende sélo Erigeron
hyssopifolius del norte de Norte América. Erigeron sect. he ale se restringe a las 14
especies del nem Jilin ie ae tienen una distribuci6n Californiana. La
especie mexicana E. ex7/is se transfiere de la sect. Cincinnactis a la sect. Lonp ocanles. Evigeron
rbizomatus ee México) y E. pee en (Guia y Durango, México) se consolidan
como Erigeron sect. sills NIX FNSOe & a eNEs, sect. nov. Las dos especies se distinguen
de g por sus famas n hojas escamosas, capitulos grandes solitarios

—

y eas largas estrechamente oblongas. La Sect. rsa nix es el elemento filogenético
mas primitivo de Erigeron y de la subtribu Conyzinae

Upon consideration of the morphology and systematics of North American
Erigeron species, we observe that modiflcations in sectional alignments should
be made. Changes discussed here deal primarily with species regarded as
“peripheral to sect. Linearifolii” in the treatment by Nesom (1992).

1. The placement of Erigeron byssoptfolius
Cronquist (1947) included Erigeron hyssopifolius Michx. within the E. foliosus

Sipa 18(4): 1161-1165. 1999

1162 Sipa 18(4)
group (sect. Pycnophyllum sensu stricto; sect. Linearifolii sensu lato, includ
ing E. hyssopifolins) but noted (p. 277) that “it certainly is not approached
by anything else in the section.” Nesom (1989) maintained E. Ayssopifolius
within the group (as sect. Linearifolzi) but later (1992) viewed it as periph-
eral co species of that group. With additional perspective on the genus, the
disparity between E. hyssopifolius and the E. foliosus group suggests that it is
more realistic to recognize the former at sectional rank, coordinate with
the species of the E. fo/iosus group in its strict sense.

Erigeron hyssopifolius differs from E. foliosus and related species in several
features, including cauline leaves abruptly reduced in size below the pe-
duncles, short, axillary, leafy shoots often produced, heads solitary on brac-
teate peduncles 5—10 cm above the level of the leaves, phyllaries 3-nerved
and all nearly equal in length, and lamina of the ray corollas not coiling.
Further, sect. Pycnophyl/lam sensu stricto is almost completely restricted to
areas along the Paciflc coast of the U.S.A. and Baja California, while E. byssopifolins
ranges across northern North America (mostly Canada) from Newfound-
land, Nova Scotia, and various New England states to Yukon and Alberta.

The nomenclature for the two sections concerned is now as follows.
ee Losec sures (G. Don) Nesom, Phyctologia 67: 79. 1989.

\yrer L. sect. Linearifolii G. Don in Loudon, Hort. Brit. 346. 1830. Lecroypr (Sundberg
nee 1987): Aster eraminifolins Pursh [= Erigeron by ues Michx. }

—_—

Species included: Erigeron hyssopifolins Michx
Erigeron L. sect. Pyenophyllum Crongq., Brittonia 6:141. 1947. Type: Eri-
veron foliosus Nutt.

Species included: the 14 species of the E. folzosus group (sensu Nesom 1992).

2. The placement of Er/geron exilis
Erigeron exilis, a species of the Mexican states of Jalisco and Nayarit, was
previously included within sect. C/neinnactis Nesom (Nesom 1989), but its
greater similarity to the species of sect. Lamprocaiules (Nesom 1994a) has
been recognized in recent study and review of Mexican Erigeron. The other
species of sect. Lamprocailes (discussed as possible relatives of sect. Linearifolii
by Nesom 1992) also are restricted to Mexico but occur in the northeastern
states of Coahuila, Nuevo Leon, and a Manne NS compared to the more
southwestern distribution of FE. exs/7

The section, as newly amended, is as follows.
Erigeron L. sect. Lamprocaules Nesom, Phytologia 76:99. 1994. Tyr: Erigeron

scopartoides Nesom

Species included: E. chiangii Nesom, E. exilis Gray ex 8. Wats., E. pattersonii

Nesom, and E. scoparioides Nesom.

These plants are perennials with stems simple or few-branched, slender

Nesom AND Noyes, Sectional delimitations in Erigeron 1163

to wiry, shiny-textured, leaves all cauline, shiny-textured, narrow, and relatively
even-sized, buds erect, and heads relatively small. The scale-leaved rhizomes
of E. exi/is are similar to those of E. chiangii and E. scoparioides; the thick,
woody, non-rhizomatous base of the gypsum endemic, E. pattersonii, prob-
ably is specialized within the section. The stipitate-glandular stems and
involucre of E. exz/is are similar to those of E. chiangii.

The ray flowers with coiling lamina of Evrigeron exilis are unusual in sect.
Lamprocaules, because the rays in other three species apparently do not coil.
Still, the overall similarity among E. exz/is and other species suggests that
they are closely related. Non-coiling (and non-reflexing) rays are uncom-
mon in Erigeron and may represent a shared specialized state in the north-
eastern species of this group.

3. The status of E. rhizomatus and E. lepidopodus
Evigeron rhizomatus and E. lepidopodus were placed by Nesom (1989) as members
of the E. folzosus group (Erigeron sect. Linearifolii sensu lato), where their
relationship as sister species was noted. Erigeron lepidopodus occurs from central
Chihuahua to northern Durango, Mexico; E. rh/zomatus 1s endemic to Catron
and McKinley counties, New Mexico, where it is considered rare and en-
dangered (U.S. Fish and Wildlife Service 1988; Sivinski and Lightfoot 1995).
In a detailed treatment of the E. /olvosus group (Nesom 1992, p. 205), un-
der the heading “Species peripheral to sect. Linearifoli7,” it again was ob-
served that these two species are closely similar in geographic range and
morphology and almost certainly related as sister species. Plants of both
produce stems from rhizomelike caudex branches, usually without clustered
basal leaves (similar to plants of the E. folzosws group), and both species pro-
duce large, solitary heads erect in bud, long, non-coiling ray corollas, and
long cypselas, unlike plants of the E. folzosws group. Cronquist (1947, p
275) observed that “Although clearly belonging to the small section Wyomingia,
{E. rhizomatus} does not seem closely related to any other known species. Its
subglabrous leaves and involucres separate it from anything else in the sec-
tion, and its peculiar habit is unique in the genus.” Indeed, of the charac-
ters noted by Cronquist to link E. rhzzomatus to sect. Wyomingia (1.e., im-
bricate involucral bracts, 4-14-nerved cypselas), the cypsela morphology
apparently was emphasized; they have little else in common. Erigeron rhizomatus
and E. lepidopodus warrant formal taxonomic recognition as a distinct and
discrete group.
Erigeron L. sect. Geronpternix Nesom & Noyes, sect. nov. Type species: Erigeron
rhizomatus Crong
Ramis caudicis longis squamifoliatis rhizomiformibus, capitulis grandibus solitariis in

alabastro erectis, ligulis non circinnatis, et acheniis longis anguste oblongis distinctus.

Perennial herbs arising from thick, fibrous roots and long, slender, de-

1164 Stipa 18(4)

cumbent, often buried, rhizomiform, scale-leaved caudex branches. Stems
erect, 6-45 cm tall, simple or with 1-3 short branches on the upper half,
stipitate-glandular at least on the peduncles, otherwise sparsely pubescent
with appressed to spreading or deflexed hairs, bearing ascending, linear to
narrowly oblong leaves. Heads large, 12—20 mm wide; ray flowers 14-45,
the lamina 6-14 mm long, white or blue-tinged with an abaxial lilac midstripe,
apparently neither coiling nor reflexing at maturity. Cypselas narrowly oblong,
3.54.5 mm long, slightly compressed radially and 2- or 4-nerved (E. /epidopodus)
or subterete and 5—6-nerved (E. rhizomatus), densely strigose to sericeous;
pappus a single series of 25—50 bristles of unequal length, with a few outer
setae. Flowering vernal. Base chromosome number, x = 9. For E. lepidopodus:
chromosome number, 2n = 18 (Ward & Spellenberg 1988); illustration (Nesom
1981). For E. rbizomatus: chromosome number, 2n = 18 (Ward & Spellenberg
1986); photographs of habit, heads, and habitat (New Mexico Native Plants
Protection Advisory Committee 1984; Heil 1995). The name is from Greek,
geron (old) and prernix (stem of a plant), alluding to the name Erigeron, to
the peculiar stems, and to the phylogenetically basal position (lower or “old
stem”) of this group.

Species included: E. /epidopodus (B. Rob. & Fern.) Nesom and E, rhizomatis
Cronq.

Recent studies by Noyes (1999) show that E. rbzzomatus and E. lepidopodus
together apparently form the most basal phyletic element within Erigeron
as well as within the entire subtribe Conyzinae. All of the species at more
basal levels of the subtribe are Er/geron, as are most of the terminal species.
Interpolated at various points in the subtribal topology, however, are other
generic-level taxa, including Conyza, which apparently is biphyletic, and a
group of South American genera (Leptostelma, Apopyros, Nefa, and Hysterionica)
recently recognized by Nesom (1994b), as well as the long-accepted and
morphologically divergent genus Aphanostephus. The taxonomic implica-
tions of the Noyes analysis are complex, but morphological differences among
various phyletically divergent infrageneric groups of species traditionally
identified as Erzgeron are in most cases so subtle that attempts to recognize
segregate genera would be virtually impossible in actual practice. The two
species of sect. Geronpternix are relatively easily distinguished as a group,

and they are justifiably recognized at sectional rank.
ACKNOWLEDGMENTS
We appreciate review comments from Richard Spellenberg and John Strother.
REFERENCES

Cronquist, A. 1947. A revision of the North American species of Erigeron, north of Mexico.
Brittonia 6:121—302.

Nesom AND Noyes, Sectional delimitations in Erigeron 1165

Hem, K.D. 1995. The Farmington District [New Mexico]. Endangered, Threatened and
Seuteve Plant Field Guide. Ecosphere Environmental Services, Inc. Farmington, NM

Nesom, G.L. 1989. Infrageneric taxonomy of New World Erigeron (Compositae: Astereae).
Phytologia 67:67-93.

Nesom, G.L. — Revision of Ergeron secc. Linearifolii (Asteraceae: Astereae). Phytologia
72:157-20

Nesom, G.L. a fa. Erigeron paler SONI ees Astereae), a new species from Nuevo

eon, Mexico. Phytologia —l
NESoM, G.L. 1994b. Subtribal ee of the Astereae (Asteraceae). Phytologia 76:193—

NEw Mexico Native PLANTS PROTECTION ADVISORY COMMITTEE. nee A handbook of rare
ndemic plants of New Mexico. Univ. of New Mexico Press, Albuquerque.

No vvES, R.D. 1999. Boge and evolutionary insights on Erigeron and allies (Asteraceae)
from ITS sequence data. Pl. Syst. Evol. (In press

SIVINSKI, R.C. and K. oe rFoot (eds.). 1995. Inventory of rare and endangered plants of
New Mexico. New Mexico and Resources rae aa Division. Energy, Min-
erals and Natural Resources Department. Misc. Pub. No. <

SUNDBERG, S.D. and A.G. JONES. 1987. cree s Hortus es (1830): An early source
of sectional names, necessitating nomenclatural changes in many genera—Aster: A case

n point. Taxon 36:97—-98.

U, : FisH AND WILDLIFE SERVICE. 1988. Zuni fleabane (Erigeron rhizomatus) recovery plan.

U.S. Fish and Wildlife Service, New Mexico Ecological Services Field Office, Albuquer-

Warb, D.E. and R.W. SPELLENBERG. hie a Chromosome counts of angiosperms of western
North America. Phytologia 61:1 2k

Warb, D.E. and R.W. ae ae Chromosome counts of angiosperms from New
Mexico and adjacent areas. Phytologia 64:390—398

1166 Stipa 18(4)
BOOKS RECEIVED

Joe LicGio and ANN Orvo Licco, 1999, Wild Orchids of Texas. Photo-
graphs by Joe Liggio; Scientific Advisior, David H. Riskind. ISBN
()-292-74712-8, hbk.). The University of Texas Press, P.O. Box 7819,
Austin, TX 78713-7819, U.S.A. $29.95. 228 pp., 87 color photos,
distribution maps.

“This beaucifully illustrated book presents all [known] 54 wild orchids of Texas.” Con-
tents include: Preface, Acknowledgments, Texas Treasures: Fifty-four Types of Orchids,
An Infinite Variety, The ae rane: Orchid, The Rael Regions of Texas, How Texas
Orchid Habitats are Described in This Book, Orchids of the Bogs and Savannahs, Orchids
of Open Sunny Habitats, Orc hids of Forests and Woodlands, Orchids of the Mountains
and Canyons, Texas Orchids by Flower Color, Texas Orchids by Genus and Species, Ap-
pendix A. Sources of Scientific Names, Appendix B. Excluded Species, Appendix C. Spe-
cies Distribution by County, Literature Cited, and Index. A full review is forthcoming in
the next issue.

WakREN L. WAGNER and V.A. FUNK (Editors). 1995. Hawaiian Biogeog-
raphy: Evolution on a Hot Spot Archipelago. (ISBN 1-56098-463-
5, pbk.). Smithsonian Institution, 470 LEnfant Plaza, Suite 7100,
Washington, D.C. 20560 (Marketing Dept. 202-287-3738). $45.00
hbk, $25.00 pbk. 467 pp., numerous

—

igures.

“Hawaiian Bhd ae compares the pIdErCe rs pie patters of many of the archipelago’s

sp

oO

with
Its ne eae history. Caan apy ve for the first time, a consistent phyloge-
netic methodology, using modern cladistic techniques, to a variety of

's—various flowering plants, birds, spiders, and insects, including fruit flies

ineages to identify
common or discordant evolutionary and biogeographic patters among the constituent spe-
cies.” Contents include: Preface, Acknowledgments, Contributors, Introduction, Geology
and Biogeography of the Hawaiian Islands, Cladistic Methods, Biogeographic Patterns of
Two Independent Hawaiian Cricket Radiations (Lampala and Peupunieollan: Chromo-
some and Male Genitalia of Hawaiian Drosophila: Tools for Interpreting Phylogeny and
Geography, Mo d

—

ecular Approaches to Biogeographic Analysis of Hawatian Drosophilidae,
Evolution of Sarona (Heteroptera, Miridae): Speciation of Geographic and Ecological Is-
lands, Comparison of Speciation Mechanisms in Web-Building and Non-Web-Building
Groups within a Lineage of spiders, Evolutionary Relationships of the Hawaiian Honey-
creepers (Aves, Drepanidinae), Biogeography of Seven Ancient Hawaiian Plant Lineages,
Wem Adaptive Radiation, and Biogeography of Hawaiian Tetramolopinm (Asteraceae,

Astereac), Phylogeny and Biogeography in Schredea and Alsinidendron (C ein oe
Historicé Biogeography and Ecology of the Hawaiian Silversword Alliance (Asteraceae):

New Molecular Phylogenetic Perspectives, Molecular Ealick ee Radiation, and
Geographic Speciation in Cyne (Campanulaceae, Lobelioideae), Patterns ance: and
Biogeography in Clermontia (Campanulaceae, Lobelioideae), Phylogenetic Analy

s of Ha-
waiian and Other Pacific Species of Scaevola (Goodeniaceae), Biogeographic a in the
Hawaiian Islands, Postscript, Literature Cited, and Index.

Sipa 18(4): 1166. 1999

NOVELTIES IN THE MYRSINACEAE
FROM THE VENEZUELAN GUAYANA
JOHN J. PIPOLY I]

Botanical Research Institute of Texas

509 Pecan Street, ae TX on 4060 U.S.A.
Jpipoly @brit.org; clusia @latinmail.com

JON M. RICKETSON

Missouri Botanical Garden

P.O. Box 299, St. Louis, MO 63166-0299, U.S.A.

)

M pa Ss hbnr
JON VICKELSON MLE OF. OV R

ABSTRACT

Preparation of a taxonomic treatment of the Myrsinaceae for the Flora of the Venezuelan
li y of anew species, Cybianthus liesneri Pipoly & Ricketson,

Guayana project resulted i
which is described, illustrated, and its systematic relationships discussed. A key to distin-
guish Cybianthus liesneri from the other species of Cybianthus subgenus Weigeltia of the Venezuelan
Guayana is provided. In addition, Parathesis viridis Lundell is found to be a member of the
genus Stylogyne. The combination, Stylogyne viridis (Lundell) Ricketson & Pipoly is pro-
posed, the species is ne illustra - and an updated description is provided. In addition,
a key to the species of Sty/ogyne found in the Venezuelan Guayana 1s presented.

RESUMEN

Estudios para preparar un tratamiento taxonémico de la familia Myrsinaceae para la Flora
of the Venezuelan Guayana dieron como resultado el descubrimiento de una especie nueva
para la ciencia. Se describe, se iluscra y se discute el parentesco de la nueva especie, Cybianthus
liesneri Pipoly & Ricketson. Se ofrece una clave taxonémica para separar Cybranthus liesnert
de las otras especies pertenecientes al género Cyhianthus subg. Weigelsia dentro de la Guayana
venezolana. Ademas, se encontré6 que Parathesis viridis Lundell se ubica mejor dentro del
género Stylogyne. Se propone la nueva combinacién, Stylogyne viridis (Lundell) Ricketson
& Pipoly, se iluscra la especie, y se discute su parentesco. Se aporta una clave para distinguir
las especies del género Sty/ogyne en la Guayana venezolana.

NOTES ON CYBIANTHUS

Of the 56 species known to occur in the area covered by the Flora of the
Venezuelan Guayana (Steyermark et al. 1995), 42 taxa, comprising 39 spe-
cies, belong to the genus Cybranthus Mart. In studying the specimens col-
lected from the region, we noted that one represented an entity we had not
seen previously. Several characters, including the abruptly basally swollen
petioles, pinnate panicles, stems with appressed brownish furfuraceous lepidote
scales and prominently long black punctate-lineations, and 4-lobed calyx

—

Sipa 18(4): 1167-1174. 1999

1168 Stipa 18(4)

placed the new species in Cybianthus subgenus Wergeltia (A. DC.) G. Agost.,
a group consisting of approximately 46 species in South America and the
Caribbean (Pipoly 1998). In the Guayana of Venezuela, subgenus Wezgeltia
is represented by six species, separable in the following key.

KEY TO CYBIANTHUS SUBGENUS WEIGELTIA
IN THE VENEZUELAN GUAYANA

7.5) em wide.

A
—~

1. Leaves narrowly oblong to oblanceolate (3.1—)4—
2. Petiole subterete; leaf blades not black aes lineare below; staminate
pedicels ca. 1 mm long; pistillate pedicels 0.5—0.6 mm long. ..... C. longifolius Miq.
. Petiole canaliculate to base; leaf blades es black punctate-lineate
below; : staminate pedicels ca. 1.5—2.5 mm long; pistillate pedicels (0.7)

1.0 ong . Surinamensis (Spreng.) G. Agost.
ee elliptic to obovate, (6.3—)8—LO(—-14) cm wide.
. Leaves membranaceous, prominently black punctate or conspicuously black

Sei ee below, base acuminate or tapering Rees ly, decurrent

on petiole to base; petiole deeply canaliculate; pistillate pedicels thin, 2—

) long.

4. Branchlets terete, not brittle, not semi-succulenc, with numerous, minute,
appressed reddish lepidote scales, inconspicuously black punctate-lineate;

leaves conspicously black punctate-lineate below, the quaternary veins

not visible; petioles 3—4.5 cm long, not decurrent. .......... C. multicostatus Mig.

at

. Branchlets angulate, brittle, semi- eae with scattered, minute,
appressed, brownish lepidote scales, densely and prominently long black
punctate-lineate, leaves densely and pee black punctate be-
low, the punctations one areole, formed by prominently raised quaternary

veins; petioles 1.5—2 cm long, decurrent on the branchlet onto rounded
pee on ie C. hesneri Pipoly & Ricketson, sp. nov.
eaves C aceous to subcoriaceous or thic a coriaceous to cartilaginous,

cree ier punctate below, base acute, barely decurrent on peti-
ole; petiole subterete with a narrow, shallow channel barely discernible
above; pistillate pedicels obsolete to thicker than long and subobsolete.
5. Branchlets angulate, with few, rounded angles below decurrent peti-
ole bases; petioles 1.3—7 cm long, decurrent onto rounded angles on
branchlets; leaves thickly coriaceous to cartilaginous; pistillate calyx
lobes deltate, apically acute; staminate calyx lobes linear-lanceolate, apically

narrowly acute to attenuate. C. grandifolius a G. Agost.
. Branchlets terete, with numerous, Taised narrow longitudinal ridge
£ sopeue l | 1.7-)2—3 cm long not a.
ne re hlec; | ‘| ‘cous to subcoriaceous; pistillate calyx lobes
very widely ovate to oblate, apically subacute to obtuse; staminate calyx
lobes ovate, apically obtuse with a small acumen at tip. ......00..... C. potiaei (Mez)
x. Agost.

The new species is described herewith.

Cybianthus liesneri Pipoly & Ricketson, sp. nov. (Fig. 1). Typr: VENEZU-
ELA. AMAZONAS: Departamento Rio Negro; Cerro de la Neblina Camp V; valley N

Prpoty AND RICKETSON, Myrsinaceae from Venezuelan Guayana 1169

of base of Pico Cardona, 00° 49! N, 66° 00' W, 1,250 m, 21-24 Mar 1984 (fr), R
Liesner & B. Stannard 16866 (WoL_otryPE: VEN; isotypes: K, MO).

Quoad folia membranacea, subter manifeste atro-punctata atque atro-punctato- lineata,

ad bases gradate decrescens, ad bases petiolaris decurrentes, petiolos profunde canaliculatos,
peaiceligs pisuiletos Braces) a= aos mm m longo Ge multicostatam valde arcte affinis, sed ab ea

{,

semi-succulentis (nec non-succulentis),
sparse deminuteque adprese brune (nec rubigni) sananats lepidotis indutis, necnon dense
atro-punctato-lineatis, foliis subter dense manifesteque

5—2 (non

tro-pun m punctis uno in ques jue areola (non lineatis), denique petiolis
4.5) mm longis, ad ramulis decurrentis (nec non-decurrentis) statim distinguitur.

Shrub or small tree to 4 m tall. Branchlets angulate, brittle, semisucculent,
7—10 mm diam., apically with scattered, minute, appressed brownish lepi-
dote scales, densely and prominently long-black punctate-lineate, glabres-
cent. Leaves pseudoverticillate; blades membranaceous, obovate, 30—43 cm
long, 10-14 cm wide, apically acuminate, the acumen 8-10 mm long, gradually
tapering to a cuneate base, decurrent to base of petiole; midrib slightly raised
above, prominently raised below, the principal secondary veins arcuate from
the midrib toward apex, prominulous above, prominently raised below, 13—
19 pairs, the tertiary veins prominulous below, perpendicular to the sec-
ondary ones, prominently black punctate below, the punctations no more
than one per each areole formed by the prominently raised quaternary ve-
nation, with scattered conspicuous (but not prominently raised) black punctate-
lineations; the margin entire, glabrous; petioles deeply canaliculate, 1.5—2
cm long, decurrent onto the rounded ridges of the branchlet, glabrous. Staminate
inflorescence: unknown. Pistillate inflorescence a columnar, pinnate panicle 10—
19 cm long, to 6 cm wide, the peduncle, rachis and pedicels densely rufous
papillate; inflorescence bract unknown; peduncle 1-2 cm long; pedicels
cylindrical, thin, 2—2.5 mm long; floral bracts unknown. Pzstzllate flowers
unknown; fruiting calyx cotyliform, spreading, 0.8-1 mm long, the tube
ca. 0.1 mm long, the lobes ovate, 0.7-0.9 mm long, 0.5—0.7 mm wide,
apex obtuse to rounded, bearing 3—5 prominently raised black punctations
medially, the margins irregular, erose to erose-dentate, glabrous. Frat (im-
mature) 2.5—3.5 mm diam., scattered translucent glandular-lepidote, the
style persistent, the stigma subcapitate, 4-lobed

Distribution and ecology.—Known only from the type, growing at 1,200—
1,300 m elevation, Cybianthus liesneri occurs in premontane wet forest, dominated
by Irzartea (Arecaceae) and several Burseraceae species.

Etymology.—This species is dedicated to Ronald L. Liesner, of the Mis-
souri Botanical Garden, prodigious collector of Amazonian plants.

Paratrypes. WENEZUELA. Amazonas: Departamento Rio Negro, trail S from Cerro

eblina Camp V, 00° 49'N, 66° 00! W, 1,200-1,300 m, 12 Apr 1984 (fr), A. Gentry G B.
Stein 46352 (MO 2-sheets, VEN n.v.)

L170 Stipa 18(4)

—

Fic. 1. Cybianthus liesneri Pipoly & Ricketson. A. Flowering branchlet, habit. B. close up of
abaxial leaf surface, showing prominent black punctations, one per areole. C. Inflorescence
branch close up, showing dense papillae. D. Close up of fruit, calyx and medicel. showing
the ovate calyx lobes, persistent subcapitate, 4-lobed stigma. A—D, drawn from type

PipOLy AND RICKETSON, Myrsinaceae from Venezuelan Guayana Lil

The following characters: chartaceous to subcoriaceous leaf blades that
are obscurely pellucid punctate below with acute bases; subterete petiole
with a narrow, shallow channel barely discernible above, and the flowers
that are either sessile or with short, stout pedicels, all indicate that C. /zesneri
is most closely related to C. mu/ticostatus. However, the angulate, brittle,
semi-succulent branchlets, with scattered minute, appressed, brownish lepidote
scales, and dense and prominent black punctate-lineations, leaves black punctate
below with one punctation per areole, shorter petioles decurrent on the rounded
ridges of the branchlet, all serve to distinguish C. /zesneri from C. multicostatus.

NOTES ON STYLOGYNE

While examining Parathesis viridis Lundell, described from the Guayana
region, we suspected that it was, in fact, a member of the genus Sty/ogyne A.
DC. The species was previously known only from fruit and some poorly
preserved flower buds. A new collection provided us with new characters,
such as the contorted corolla,

—

inear-lanceolate anthers with sagittate bases
and longitudinal dehiscence, long style with punctiform stigma, leaving
no doubt that Parathesis viridis should be transferred to Stylogyne. There-
fore, we propose the new combination here and include the new data gleaned
from the second collection known for the species. A key to the species of
the genus in the Venezuelan Guayana is provided first, below.

KEY TO SPECIES OF STYLOGYNE IN THE VENEZUELAN GUAYANA

1. Inflorescence, peduncle, rachis and/or ae ae! to densely papillose,
of simple to few-celled papillae, often obsc

—

2. Anthers narrowly lanceolate, reddish-con ie rous when dried; me
opaque; ovary minutely rufous puberulent apically viridis (Lundell)

Ricketson & ee comb. nov.
2. Anthers oblong, whitish-yellow when dried; inflorescence translucent pink

or pinkish-red; ovary glabrous apically ..............0.:000 S. orinocensis (Kunth) Mez

1. Inflorescence, peduncle, rachis and/or pedicels glabrous.
3. Leaf margins conspicuously crenate; growing in gallery forests on slc
near streams, ca. SOO m ha S. lasseri (Lundell) Pipoly
3. Leaf margins entire, obscurely eee in wet or swampy
lowland forests, 30-450(— 800) m eleva
4, Mature fruits 10-14 mm diam. ee -2 slobase: leaf blades chartaceous
to coriaceous, cusaany nitid aboveS. atra Mez
4, Mature fruits 4—7(—9) mm diam., globose; leaf blades penne 1S
usually at aa S. micrantha (ant ) Mez

The transfer and new description is included herewith.

Stylogyne viridis (Lundell) Ricketson & Pipoly, comb. nov. ue a Parathesis
viridis Lundell, Phytologia 56:26. 1984. Type. VENEZUELA. ou MAZONAS: 0-1 km
So s de Rio Negro, 01° 51' N, 67° 03' W, 120 m, a ep R.

ae 9046 (HOLOTYPE: MO; isoTyPE: VEN n.v.).

1172 Sipa 18(4)

Fic. 2. Stylogyne viridis (Lundell) Ricketson & Pipoly. A. Flowering branchlet habit, show-
ing wings, and leaf punctations. B. Close up of abaxial leaf survce, showing punctations
and punctate-lineations. C. Corymb detail, showing reflexed corollas and persistent styles.

Corolla and adroecium, separated from calyx and developing fruit. E. Stamen, adaxial
view, showing hastate

—

dase and longitudinal slits. F Stamen, lateral view. G. Stamen, abaxial
view, showing darkened connective. H. Pedicel, calyx and young fruit, showing minute
rufous puberulent ovary summit. A—H, drawn from R. Liesner 4182.

PIPOLY AND RICKETSON, Myrsinaceae from Venezuelan Guayana 1173

Tree to 4 m tall. Branchlets 2-2.5 mm diam., angulate, with ridges form-
ing “wings” ca. 3 mm high, spiralled around the stem to follow the mar-
gins of the decurrent petiole bases, glabrous. Leaves alternate, chartaceous,
lanceolate to elliptic, 14-19 cm long, 4.04.7 cm wide, apically long acuminate-
attenuate, the acumen 1—1.8 cm long, basally acute to obtuse, midrib prominulous
apically, then canaliculate basally above, prominently raised below, the secondary
veins 20—24 pairs, prominulous above and below, connected by a submar-
ginal looping vein ca. 4 mm from margin, with periodic secondary veins
extending toa second minor submarginal connecting vein 1 mm from margin,
minutely scrobiculate above, densely and prominently orange punctate and
punctate-lineate below, the margins flat, entire, decurrent to base of peti-
ole; petiole marginate, roughly trigonal in transverse section, marginate,
6-9 mm long, glabrous. Bisexual inflorescence a terminal, pyramidal, bipin-
nate panicle 5.5—8 cm long, 6—7.5 cm wide; inflorescence bract unknown;
peduncle 1.1—1.5 cm long; floral bracts coriaceous, minute, linear, 0.4—0. 6
mm long, 0.1—-0.2 mm wide, apex rounded, densely orange punctate, the
margin glandular-ciliate, early caducous; pedicels 4-6 mm long, minutely
glandular-papillate. Bisexwal flowers chartaceous: calyx campanulate, the sepals
free, ovate, 1.1—1.3 mm long, 0.7—-0.9 mm wide, apex rounded to obtuse,
the margins irregular, somewhat erose apically, very sparsely and minutely
glandular-ciliolate; corolla chartaceous, very openly rotate, 4.4—5 mm long,
the tube 0.9-1.1 mm the lobes reflexed distally and rolled at anthesis, ob-
long, 3.5—3.9 mm long, 1.3—1.5 mm wide, apex very broadly rounded toa
short acumen, densely and prominently orange punctate and punctate-lineate,
the margin entire, opaque, glabrous; stamens 3.3—3.6 mm long, the fila-
ments flat, 1.8-2 mm long, adnate to the corolla tube 0.9—1.1 mm, the
apically free portion ca. 0.9 mm long, the anthers concolorous, linear-lan-
ceolate, 2—2.3 mm long, 0.3—0.5 mm wide basally, apically attenuate, ba-
sally deeply sagittate, dehiscent by wide longitudinal slits, the connective

ark brown; pistil obturbinate, 4.7-4.9 mm long, the ovary obovoid, 2—
2.2 mm long, 1.5—1.7 mm diam. toward apex, narrower below, apically
rufous puberulent, the style thin, 2.6-2.8 mm long, the stigma puncti-
orm. Frazt unknown.

Distribution.—Endemic along the Rio Negro, south of San Carlos de Rio
Negro. It grows at 120 m in elevation. No collections are known from the
Colombian side of the river, however it should be expected.

Ecology and conservation status.—Seasonally flooded primary forests and secondary
areas. Because of its restricted distribution, it should be considered threatened.

Etymology.—The specific epithet comes from the latin meaning “green,”
presumably because of the leaf color.

cimens examined. VENEZUE S of Carlos de Rio Negro and
ee river for 2 km, 01° 55'N, ra W, 120 m, 4 Dec 1977 a fr), R. Liesner 4182 (MO).

1174 Sipa 18(4)

The minute glandular-papillae of the pedicels makes it most easy to confuse
Stylogyne viridis with S. orinocensis. However, the concolorous, linear-lanceolate
anthers, opaque inflorescence rachis and ovary minutely glandular-puberu-
lent apically all easily distinguish Stylogyne viridis from S. orinocensis.

ACKNOWLEDGMENTS
This paper is a result of the Flora of the Venezuelan Guayana project at
the Missouri Botanical Garden, under the auspices of the Julian A. Steyermark

Fund and NSF grants BSR-8717303. BSR 9045532, and BSR 9201044.
The illustrations were prepared by Jon Ricketson.

REFERENCES
Pipoty, J. 1998. The genus Cybianthus in Ecuador and Peru. Sida 18:1—160.

STEYERMARK, J., P. Berry, and B. Hoist (eds). 1995. Flora of the Venezuelan Guayana. Vol.
lL. Introduction. Missourt Botanical Garden and Timber Press. Pp. 320.

NEW SPECIES OF CLUSIA (CLUSIACEAE)
FROM THE CORDILLERA OCCIDENTAL
OF COLOMBIA

JOHN J. PIPOLY III
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX TOIVE ue U.S.A.
ppipoly @brit.org; clusia@lat l com

ALVARO COGOLLO P.

Fundacién Jardin Botdnico, Joaquin Antonio Uribe
Apartado Aéreo 51407, Medellin, Antioquia, COLOMBIA
acogollo@latinmail.com; jardinbo@ md. impsat. net.co

ABSTRACT

Fieldwork conducted during a survey of Las Orquideas National Park in Antioquia, Colombia,
esulted in the discovery of two undescribed species of Clusia section Anandrogyne. Clusia
ennai Pipoly & Cogollo and C. paisarum Pipoly & Cogollo are described and illustrated,
and their respective distributions, oe conservation status, etymology of epithets and

phylogenetic relationships are elucidate

RESUMEN

El trabajo de campo durante una investigacion de la flora del Parque Nacional Natural
‘Las Orquideas,” ubicado en el Departamento de Antioquia, Colombia, dio como resultado
ms desc pe ee de dos especies nuevas, pe Trenecicntes al género Cues secci6n A nandrogyne

ray

il istrac 1ones

senlont y Salone en cuanto a la conservaci6n, asf como la tiers de ic epitctos
especificos y el parentesco de Clusia deminuta Pipoly & Cogollo y de C. patsarum Pipoly &
Cogollo
The genus Clusia section Anandrogyne Planch. and Triana is now known
to contain more than 70 species (Pipoly 1995, 1998) and is defined by the
largely anantherous staminodes of the pistillate flowers and the pluriseriate,
acropetally longer stamens of the staminate flowers, the latter with anthers
dehiscent by wide longitudinal slits. While carrying out fieldwork to document
the plant diversity of Las Orquideas National Park, two new species were
encountered and are described herewith.
Clusia (§ Anandrogyne) deminuta Pipoly, sp. nov. (Fig. 1). Typ. COLOM-
NTIOQUIA: Mpio. Urrao; Corregimiento La Encarnacién; Trail to Parque Nacional

Natural “Las Orquideas,” first hill between Quebrada el Aguacate and Quebrada San

Sipa 18(4): 1175-1182. 1999

1176 SIDA 18(4)
José, 06° 27'N, 76° 13' W, 2,200—2,400 m, 8 Feb 1995 (fr), J. Pepoly, J. Ramirez G
J. Arias 18595 (HoLoryPe: JAUM; tsorypes: BRIT, FMB

Quoad panic ulam 5-floridam, folia subsessiles vel sessilies, petiolos late marginatos sepala

de arcte affinis sed ab ea ramulis tetragonis (non teretibus),

ines 6, C. pan val
ndatis),

lami ‘| loratis (non ellipticis) ad apices acuminatis (nec rotun
pec dicellis tetragonis 2 $32 mm longis (non cbeolotic): necnon habictu epiphytico (non

terrestri) pert facile cognoscitur.
Epiphytic shrub to 3m tall; latex white. Branchlets tetragonal, with the
angles formed by small rounded ridges running from the side of each peti-
3mm diam. between nodes,

<

ole base to the center of the petiole above it, 2.5—
3—4 mm diam. at the nodes; semisucculent, the bark smooth, shiny, exfoli-
elabrous. Leaves subsessile to very short petiolate; blades thinly co-
1.5 cm long, (1.2-)2.0-2.4

ating, g
riaceous, linear, oblong or lorate, (S.2—)6.0-1
cm wide, apically acuminate, the acumen 4—8 mm long, basally broadly
rounded, the midrib prominently raised above and below, the secondary
veins numerous, perpendicular to the midrib, prominently raised above and
below, the submarginal collecting vein perpendicular to the secondarie
ioe raised above and below, the tertiary veins not visible from above,
prominulous below, glabrous and dull above, pallid below, the latex canals
not obvious, cane numerous rubiginous dots below, the margin revo-
lute, decurrent on the petiole; petiole subobsolete to 4 mm long, deeply
canaliculate and widely marginate throughout, glabrous, decurrent on e1-
ther side of the stem and almost touching the petiole base of the opposite
leaf. Staminate inflorescence and flowers unknown. Pistillate inflorescence term1-
nal, dichasial, 5-flowered, with two basal lateral flowers followed by a ra-
chis segment and 3 terminal flowers; peduncle tetragonal, 1.7—2.5 cm long,
inflorescence bracts 2, qualitatively identical to foliage leaves but membranaceous
15-17 mm long, 3—3.5 mm wide, midrib raised in a sunken furrow above,
prominently raised below, the secondary venation not visible, the petioles
obsolete to 2 mm long; upper peduncle tetragonal, 9-11 mm long; floral
bracteoles 2, below the two basal, lateral flowers, stiffly coriaceous, widely
ovate, 3—3.5 mm long, 2.4—2.6 mm wide, apex broadly rounded but with
a small acuminate-mucronulate tip, ventrally (abaxially) keeled medially,

the margin opaque, entire; pedicels of basal lateral flowers like the peduncle
Z ame mm wide,

upper bracteoles as in lower bracteoles but 3—3.2 mm long
&,
3.1 mm long.

terminal flower cluster pedice’s as in peduncle except 2.8—
Pistillate flowers with sepals 4, decussate, chartaceous, the outer suborbicu-
lar, cucullate 4—4.2 mm eae. 3—3.2 mm wide, apically broadly rounded

the margin opaque, entire, inner sepals as in outer except margin irregu-
larly notched; petals 6, contorted, membranaceous, obovate-spathulate, as
in sepals except constricted basally; staminodes 5, alternate with the car-
pels (opposite carpel suture), minute, anantherous, subulate, ca. | mm long,

PIPOLY AND COGOLLO, A new species of Clusia from Colombia Ee

-————,
6mm

0.5cem

1G. 1. Clusia deminuta Pipoly. A. Fruiting branchlet. B. Close up of abaxial leaf surface,
showing revolute margin and rubiginous dots. C. Piscillate inflorescence in
section, showing sepal (left)

A

, piscil (upper center), petal (right), and pedicel with floral bracteoles
—E, drawn from type.

(lower center). E. Fruit

0.3 mm wide, early caducous; pistil 5-carpellate, oblongoid, ca. 3 mm long,
1.3 mm diam., the styles obsolete, the stigmas 5, cuneiform, brick red, the
surface appearing papillate. Mature fruit ovoid, 1.3—2 cm long, 0.8—1 cm

1178 Stipa 18(4)

diam. when dried, sutures furrowed, the styles stout, ca. 1.5 mm long, the
stigma concave, deltoid (triangular with rounded corners), the seeds small,
numerous, with orange arils.

Distribution.—Apparently endemic to Parque Nacional “Las Orquideas,’
in the Municipio of Urrao and the Corregimiento of La Encarnacién, De-
partment of Antioquia, Colombia, on the Cordillera Occidental of the Andes,
growing from 1,300—2,400 m elevation.

Ecology and conservation status, —Clusia deminuta 1s a canopy epiphyte in
the pluvial premontane, cloud and elfin forest, where it forms large indi-
viduals that often acquire a weight sufficient enough to break the branches
of the host tree. In the premontane forest, it is restricted to the tops of emergent
trees, whereas higher, in the cloud and elfin forest, it may be found in tangles
of fallen vegetation. It requires nearly open liehe so is very susceptible to
the periodic, catastrophic “blowdowns” seen on radar imagery for the re-
gion, mostly attributable to seismic activity. Even though the species has a
locally common distribution, | would still consider it threatened because
the entire region is prone to frequent and violent seismic activity. It should
be noted that the “elfin forests” inhabited by C/asia deminuta are some of a
very few true elfin forests found on the South American continent, mostly
because the mountains on which they occur are not high enough to permit
existence of subpdramo and paramo formations.

Etymology. —The epithet “deminuta” refers to the small stature of the plant,
its leaves, branchlets, flowers and fruit.

Paratyprs. COLOMBIA. Sree ese Mpio. Urrao; Corregimiento La Encarnacion, Parque
Nacional Natural “Las Orquideas,” Sector Calles, Quebrada La Bironda, 06° 31' N, 76
19' W, 1,300-1,500 m, 3 ‘Ap 1992 (fr), D. Cardenas & E, Alvarez 3261 pe JAUM,
MQ), Vereda Calles, right bank of Rfo Calles, on the mountain range NW of Cabafa Calles,

06° 32'N, 76° 19' W, 1,450 m, | Dec 1994 (fr), A. C ates hes ee FMB, JAUM),

see anent ete ine pluvial forest inventory plot, 06° 32'N, 19' W/, 1,450-1,500
1, 28 Nov 1993 (scer.), J. Pipoly, A. Cogollo et al. 17292 (BRIT, : MB, arti M); Por 1 oe
dl Parque econet Nacaral Las Orquideas,” Vereda Calles, 06° 32'N, 76° 19'W, 1,

500 m, 30 Nov 1993 (pisc. fl bud), J. Prpoly et al. 17569 Ree FMB, JAUM).

The cuneiform stigmas, appearing papillate, with 5 carpels and 5 very
reduced, subulate staminodes without antherodes, clearly place Clusia demimuta
within section Avandrogyne Planch. & Triana. Section A nandrogyne is by far
the largest and the most complex within the genus, containing 75 species
(Pipoly et al. 1995, 1998), including at least 21 as yet undescribed. The
five-flowerec | panicle, subsessile or sessile leaves, widely marginate petioles
(when these are developed), calyx of four sepals and corolla of six petals indicate
that Clusia deminita is most closely related to the vicariant C. popayanensis
Planch. & Triana. Clusia popayanensis is a poorly known species, from the
Chocé floristic region of western Cauca Department, growing in the subparamo
thicket life zone. Clusia deminuta is easily separated from C. popayanensis by

PIPOLY AND COGOLLO, A new species of Clusia ft °m Colombia 1179

its tetragonal branchlets, linear, oblong, or lorate leaf blades with acumi-
nate apices, tetragonal pedices| 2.8—3.2 mm long, and epiphytic habit.

Clusia deminuta 1s an important, locally common endemic from Las Orquideas
National Park. The elfin forest area at the Park’s northern boundary is ex-
tensive and our exploration was very lim ited. Based on collections from the
area, the northernmost areas of the Cordillera Occidental of Colombia is
home to large numbers of undescribed, vet ecologically important and very
conspicuous species of flowering plants.

ee paisarum Pipoly, sp. nov. (Fig. 2). Typr. COLOM-
ANTIOQUIA: Mpio. Urrao; Corregimier to La Encarnaci6n; Trail to P ree Nacional
— ‘Las Orquideas,” first hill peter 1 Quebrada el Aguacate and Quebrada San
José, 06° 27' N, 76° 13' W, 2,200—2,400 m, 8 Feb 1995 (fr), a es J. Ramtrez &
J. Arias 18585 (HoLotyPr: JAUM; isoryp2s: BRIT, COL, FM

Propter inflorescentiam terminalem atque flo-es congestos, lamina coriaceaque elliptica
vel oblonga, pedunculos quadratos, ovarium 7-carpellatum, denique fructus ficagan oa
rostratus, C. casstnoidi valde arcte affinis, sed ab ec ramulis reg mere teretibus), lamin
acuminatis (non rotundatis), petiolis 112 2 (non usque ad 1.0) cm longis, necnon ote
5 (non 4), atque petalis G(nec 5), statim senanas ilis.

Terrestrial shrub to 3 m tall; latex yellow. Branchlets tetragonal, the angles
formed by acute angles cease from each petiole base to the center of the
petiole 90° from and above it, 4.5—5(—7) mm diam., not swollen at the nodes;
semisucculent, the bark es sordid, not eta: glabrous. Leaves
petiolate; blades coriaceous, elliptic to narrowly oblong or rarely lanceolate,
(7.0—)11.5-15(-17) cm long, (3.2—4—5.5(—G6) cm wide, apically short acuminate,
the acumen 5—10 mm long, basally acute, midrib raised but canaliculate
above, the channel decurrent to petiole base, pronmlneacly raised below, the
secondary veins 28—34 pairs, diverging at approxmately 45° from the mid-
rib and arcuate, barely prominulous or inconspicuous above, prominulous
below, the submarginal connecting vein barely prominulous above, prominulous
below, the secondary veins alternating with shorter intersecondary veins that
terminate before reaching the connecting vein, the tertiary veins inconspicuous,
the latex canals numerous, linear, black, conspicuous below only on imma-
ture leaf blades, glabrous and dull above, pallid below, the margin entire,
flat, decurrent on the petiole; petiole deeply canaliculate and marginate
throughout, (1.1—)1.3—1.5(—2.2) cm long, glabrous. Staminate inflorescence
and flowers unknown. Pistillate see terminal, a 6-flowered, congested
cyme; peduncle tetragonal, (1.7—)4.0-5.5 cm long; inflorescence bracts 2,
qualitatively identical to the vegetative leaves except blades (2.0-)3.0-8.5
cm long, (0.7—)1.2—3 cm wide, the petioles 0.5—1.0 cm long; upper pe-
duncle tetragonal, 5-8 mm long; floral bracts 2, cartilaginous, ovate, 10—
12 mm long, 6.5—7 mm wide, apically acute to attenuate, prominently keeled,
medially strongly rugose, the margin stramineous, entire; floral bracteoles

1180 SIDA 18(4)

Fic. 2. Clusia paisarum Pipoly. A. Een a anc Bien Bb. TAH OrESCEOEE, earee al view. C. Dis-
sected pistillate flower bud, sl g ). D. Infructescence
A-D, drawn from type.

Pipoty AND CoGoLLo, A new species of Clusia from Colombia 1181

2, as in floral bracts but widely ovate, 6.8—7.2 mm long, 5,8—6.2 mm wide,
apically broadly rounded to an acute tip, prominently rugose medially, the
margin stramineous, entire; pedicels obsolete. Pisté//late flowers: sepals 5, the
outer two opposite, the inner three contorted; outer sepals stiffly eanulaet
nous, suborbicular, 8.8-9.2 mm long, 10.1-10.5 mm wide, apically very
broadly rounded, somewhat cucullate, medially somewhat rugose, the margin
opaque, entire; inner sepals acropetally larger, chartaceous, stramineous, ovate,
to 9.4 mm long, 7.3 mm wide, apex broadly rounded, the upper margin
thin, translucent, irregularly incised; petals 6, coriaceous, contorted, ob-
long, 13-14.1 mm long, 6—-7.5 mm wide, apically slightly cucullate, the
margin opaque, entire; staminodes 7, alternate with the carpels, anantherous,
connate into a small ring, oblate, 0.7 mm long, |.3—1.5 mm wide, apically
sharply acuminate; pistil 7-carpellate, obovoid, 9.8—10.2 mm long, the ovary
ca. 7-7.3 mm long, 7.3—7.5 mm wide, the styles 2-3 mm long, the stig-
mas concave, black, cuneiform, 1.3—1.5 mm long, 1.2—1.4 mm wide, smooth.
Mature fruit globose, 1.3-—2 cm long and wide, Ene carpels 7, not obvious,
without obvious suture lines, the styles thin, 2-3 mm long, giving a beaked
(rostrate) appearance to the fruit, the ti amas: as in the flowers, the seeds
small, numerous.
Distribution.—Apparently endemic to the type area near Parque Nacional
“Las Orquideas,” in the Municipio of Urrao and the Corregimiento of La
Encarnacién, Department of Antioquia, Colombia, along the western slopes of
the Cordillera Occidental of the Andes, growing from 2,200—2,400 m elevation.
Ecology and conservation status.—Clusia paisarum is a terrestrial tree grow-
ing in remnant montane pluvial forest, near small watercourses. It is lo-
cally common along the margins of remnant forest, but particularly abun-
dant on the ridges above roadcuts. It is also an important element of these
remnants because it is deeply rooted and in fact, is often used to tie pack
animals to as they rest along the paths. However, it does not attain a diam-
eter over 10 cm DBH, and perhaps for that reason, and its copious, sticky
yellow latex, it is rarely cut down. Despite significant effort, no staminate
plants were found, but there were significant numbers of trees not in flower,

some of which may have been staminate.

Etymology.—The specific epithet “paisarum” is derived from the local Spanish
adjective “Paisa,” a colloquialism for Colombians inhabiting the coffee-growing
region comprised of the Departments of Antioquia, Risaralda, Caldas and
Quindio. The region around Urrao, Antioquia is particularly noteworthy
for its production of coffee and grenadilla, a species of Passiflora.

Paratype. COLOMBIA. Antioquia: Mpio. Urrao; Corregimiento La Encarnaci6n,; Trail
to Parque Nacional Natural “Las Orq Bees first hill between Quebrada el Aguacate and
Quebrada San José, 06° 27' N, 76° 13' W, 2,200-2,400 m, 8 Feb 1995 (fr), J. Pzpoly, J.
Ramirez G J. Arias 18580 (BRIT, oe FMB, JAUM)

1182 Sipa 18(4

Ww

Clusia paisarum is most closely related to C. cassinoides Planch. & Triana,
with which it shares of its yellow latex, terminal, congested inflorescence,
coriaceous usually elliptic or oblong leaf blades, tetragonal peduncles, 7-
carpellate ovaries and globose, beaked fruits. However, Clusia paisarum can
immediately be separated from C. cassinoides by the tetragonal stems, short
acuminate leaf apices, longer petioles, more numerous sepals and petals.

This species belongs to a group of species within the large C/us/a section
Anandrogyne with yellow latex. This group includes such problematic taxa
as Clusia stenophylla Standl. and C. /ongistyla Cuatrec., the former found from
Mesoamerica to the Darien of Panama, and the latter from the northern pluvial
lowland forests of Antioquia and the Choco of Colombia, southward to
Esmeraldas, Ecuador, an area still within the Choco Floristic Province. However,
both Clusia stenophylla and C. longistyla have much larger oblong or elliptic
leaf blades with broadly rounded apices and bases, pyramidal panicles, ex-
tremely long, alate petioles (3—8 cm long), 5-carpellate fruits and long, thin,
persistent styles, clearly distinguishing them from both Clusia paisarum and
Clusia cassinoides.

ACKNOWLEDGMENTS

This study is the result of a generous grant from the National Geographic
Society, number 5575-95, to John Pipoly (BRIT-PI) and Alvaro Cogollo
(JAUM-Co-PI). We thank our respective institutions for time to carry out
the work. We also thank the staff of the Minsterio del Ambiente, regional
office in Medellin, and the staff of the Parque Nacional Natural Las Orquideas,
especially Ovideo Alvarez, for critical logistic support, and for the cordial
hospitality afforded us by the residents of La Encarnacion and vicinity. Thanks
are due also to the curator of JAUM, Biol. Juan Guillermo Ramirez, and
students of Dr. Ricardo Callejas, from the Universidad de Antioquia, espe-
cially J. Arias and G. Munoz. Luzmarina Velez of JAUM also provided critical
logistical and organization skills, contributing to the success of the study.
Lastly, the nice illustrations were provided by Linda Helms in fulfillment
of her internship in botanical illustration at the Botanical Research Insti-
tute of Texas.

REFERENCES

Pipoty, J. 1995. Cybianthus fabiolae, a new a with a synopsis of C/wsia section A nandrogyne
(Clusiaceae) in Guayana. Sida 16:737—
.. D. Kearns and P. Berry. 1998. Cla. In: P. Berry, B. Holst and K. Yatskevich,
eds. Flora of the Venezuelan Guayana. Vol. 4, Caesalpiniaceae—Ericaceac. Missouri Bo
tanical Garden Press. St. Louis, Missouri. 260-294

SARRACENIA ROSEA (SARRACENIACEAE),
A NEW SPECIES OF PITCHER PLANT FROM THE
SOUTHEASTERN UNITED STATES

ROBERT EC. NACZI and ERIC M. SOPER!
Department of Biological Sciences
Northern Kentucky University
Highland Heights, RY 41099, U.S.A.
FREDERICK W. CASE, JR. and ROBERTA B. CASE?

7275 Thornapple Lane
Saginaw, MI 48603, U.S.A.

ABSTRACT

on

Sarracenia rosea is described as a new species from the Gulf Coastal Plain of southern
Alabama, northwestern Florida, southwestern Georgia, and southeastern Mississippi. Pink

petals, large flowers, short scapes, pitchers with thick lips, and lips deeply concave in dor-
siventrally pressed pitchers are among the features that distinguish S. rosea from its closest
relative, S. purpurea. Sarracenia rosea most often grows in seeny to iphely: shaded, wet, acid
sites in pinelands. Its infrequent occurrence within asmall g

o

of concern.

RESUMEN

Se describe como nueva Sarracenia rosea de la Llanura Costera del Golfo del sur de Ala-
bama, noroeste de Florida, sudoeste de Georgia, y sudeste de Mississippi. Los pétalos rosas,
HOES earns, eer: cortos, Hepcnies con labios eee, y labios Snuy céncavos en los

1 las m j res stinguen S. vrosed

de su paricnie mas cercano, S. purpurea. ey rosea crece la mayor ca las veces en
lugares himedos dcidos en pinares abiertos, de soleados a ligeramente sombrios. El vivir en

un drea geografica pequefia, donde aparece con poca frecuencia hace que su conservaci6n
sea de interés

INTRODUCTION
The Western Hemisphere pitcher plants (Sarraceniaceae) are a small family
(15-17 species in 3 genera) of perennial herbs endemic to North America
and northern South America. Tubular, pitcher-like leaves (“pitchers”) en-
able these plants to lure, trap, digest, and absorb nutrients from a wide variety

'Current address: College of Dentistry —D155, University of Kentucky, Lexington, KY
40536, U.S.A

“Deceased 8 June 1998

Sipa 18(4): 1183-1206. 1999

1184 Stipa 18(4)

of prey, usually arthropods. Most of these pitcher plant species inhabit sunny,
wet, nutrient-poor sites such as bogs, seepages, and wet savannas. The car-
nivorous habit, peculiar morphology, and unusual ecology of these plants
have attracted much interest from horticulturists, ecologists, and system-
atists. Despite such attention, the ecology, taxonomy, and phylogeny of the
Sarraceniaceae remain poorly understood.

The largest genus in the family is Sarracenia (10 or 11 species). The most
morphologically divergent member of Sarracenia is 8. purpurea L. It is the
only species with erect hoods and, consequently, the only species whose pitchers
collect rainwater. Other Sarracenia species have hoods that cover the mouths
of the pitchers, which are moist inside, but not full of water. Sarracenia purpurea
is the only species in which the pitcher tube is curved, the widest part of
the tube is near the middle, and the style expansion is glandular-puncticulate.
Congeners have straight tubes, tubes widest at or near their mouths, and
nonglandular style expansions. Only S. purpurea and S. psittacina possess short,
decumbent pitchers with wide wings. Other species of Sarracenia have rela-

pe

tively long, erect pitchers with relatively narrow wings.

Sarracenia purpurea is often locally abundant within its vast geographic
range, which 1s the largest in the family. Its reported natural range spans
32 degrees of latitude (30-62° N) and 70 degrees of longitude (53—123°
W), from southeastern Louisiana, southern Alabama, and northern Florida
north in the souchern Appalachian mountains and along the Atlantic Coast
to eastern Newfoundland and then west across the northeastern United States
of America and much of southern Canada to northeastern British Colum-
bia (Macfarlane 1908; McDaniel 1971; Cody & Talbot 1973; Rouleau &
Lamoureux 1992).

Largely because of its distinctive morphology, relative frequency, and wide
range, S. purpurea has a four-century history of study and is the best known
member of its genus (Lloyd 1976). Authors have described several infraspecific
taxa of S. purpurea (Schnell 1979, 1993; Schnell & Determann 1997; Hanrahan
& Miller 1998). In our ongoing field, greenhouse, and herbarium studies of
the taxonomy of S. purpurea, we have discovered that one of these taxa (S.
purpurea var. burkii D.E. Schnell) is more distinct than previously recog-
nized and deserves recognition as a new species.

MATERIALS AND METHODS

We have investigated the morphology, distribution, and habitats of S.
purpurea and the new species in the field at as many sites as possible throughout
its geographic range. For the new species, we studied it at 26 sites in Ala-
bama, Florida, and Mississippi. We studied 8. purpurea in 4 Canadian prov-
inces and 12 states of the U.S.A. We also borrowed specimens from or studied

Naczi eT AL., A new Sarracenia from the southeast 1185

specimens at the following 20 herbaria: AUA, DHL, FLAS, FSU, IBE, KNK,

M U, NY, OS, PH, TENN, US, USAM, USCH,
VDB, and WKU. Abbreviations of herbaria are those of Holmgren et al.
(1990).

Previous authors vary in their terminology for the morphology of Sar-
racenta purpurea. In order to standardize terminology and make our descrip-
tions of characters unambiguous, we picture and use the following terms
for portions of the pitchers: hood, mouth, lip, tube, wing, and petiole (Fig.
1A). Terms for the description of reproductive morphology are relatively
straightforward, with the exception of style expansion, which we have il-

>

lustrated in Fig. 1B.

Of the herbarium specimens we examined, including our own collections,
we selected a representative subset to measure for statistical analyses. We
chose only mature specimens that had grown in sunny or lightly shaded
habitats because pitchers etiolate and pitcher plants exhibit reduced growth
in heavily shaded habitats. For specimens collected by others, we used habitat
information on labels and the presence of reddish venation strongly con-
trasting with the ground color of pitchers as evidence of growth in high-
light environments. We also chose specimens that exhibited the full range
of morphologic variation for 8S. purpurea and the new species and that origi-
nated from throughout the ranges of the two taxa. The set of specimens we
measured includes all of the subspecies and varieties currently recognized
in S. purpurea. Each measured specimen of the new species is denoted by an
asterisk following its herbarium of deposit in the citation of types and of
representative specimens. Citations of measured specimens of S. purpurea
can be found in the Appendix. Specimen citations have been purposely
abbreviated because of conservation concerns.

We measured at least 74 specimens for most characters of S. purpurea and
at least 39 specimens for most characters of the new species. Sample sizes
vary for the characters since some collections lacked a particular feature or
the manner of specimen preparation made measurement impossible. We
measured each character only once per specimen. When measuring a struc-
ture that is present more than once on a specimen (e.g. petals), we mea-
sured the one with the greatest value for a measurement. When measuring
the width or thickness of a structure, we measured it at its widest or thick-
est point. We measured maximum distance of the lip from the horizontal,
lip thickness, and style arm length as in Fig. 2. For style arm length and
petal length, we measured only styles and petals that were fully expanded.
To assess the degree to which S. purpurea and the new species differed from
each other for each character, we used the independent-samples ¢-test. Be-
cause the variances of several characters are heterogeneous (as determined

1186 Sipa 18(4)

Fic. 1. ie oe es of Sarracenia purpurea and S. rosea. A. Pitcher. B. Flower after shedding
of petals. The flower is turned to reveal the style expansion, which faces the ground in these
pendent flowers.

by the Bartlett chi-square test for homogeneity of group variances), we trans-
formed the data with the common logarithm before conducting the f-tests.
All statistical analyses were performed on a Macintosh computer using SYSTA’
version 5.1 (Wilkinson 1989). In Table 1, we report sample sizes, summary
statistics, and ¢-values for the measurements of the characters we discov-
ered that best distinguish S. purpurea and the new species.

To study the lectotype of S. purpurea (McDaniel 1971), which is plate 70
of Catesby (1738), we measured the plate directly as if it were a pressed
specimen. Direct measurement is justified because Catesby’s depictions of
S. purpurea and a frog on the plate are life-sized. In text accompanying plate
70, Catesby states, “These frogs are of various sizes, tho’ commonly about
the bigness of the figure. ...” We did not include measurements of the lec-
totype in the data or analyses presented in Table |

Naczi ET AL., A new Sarracenia from the southeast 1187

1. maximum distance 3. style arm length
from horizontal

2. lip thickness

A B

Fic. 2. Methods of measurement of specific diagnostic features of Sarracenia purpurea and S.
rosea. A. Distal portion of pitcher, with diagnostic features of the lip indicated. B. Flower
after shedding of petals, with style arm length indicated.

As an additional means of studying the morphology of S. purpurea and
the new species, we cultivated plants of them under common conditions in
the Case greenhouse in Saginaw, Michigan. In the greenhouse, we grew plants
from 8 populations of S. purpurea (3 from the southern Appalachians, 3 from
the Coastal Plain of North and South Carolina, and 2 from Michigan) and
7 populations of the new species (Alabama and Florida). Plants were culti-
vated for 15—25 years in pots with soil from the same source and watered
from a common supply, as described by Case and Case (1976),

For determination of the flowering period of the new species, we consid-
ered only specimens bearing at least one fully expanded petal. To determine
the geographic range of the new species, we used only herbarium specimens
that we examined; each point on the map is based on at least one herbarium
specimen.

1188 Sipa 18(4)
RESULTS AND DISCUSSION

anes oe rosea Naczi, EF. W.Case, & R.B.Case, sp. nov. (Figs. 3, 8). Tyer:
S.A. FLORIDA. Liberty Co.: SW of Telogia, eae, National eee 17 May
]. Naczi 3016 (HOLOTYPE: MICH *: isotypes: NY*, US

Sarracenia purpurea ssp. venosa (Raf.) Wherry var. burki7 D.E.Schnell, Rhodora 95:8. 1995.
Type: U.S.A. ALABAMA. Washington Co.: off U.S. 45 near Deer Park, 4 Apr 1992,
Schnell s.n. (AOLOTYPE: NCU, n.v.). Schnell’s taxonomic concept is clear from his color
photograph (Schnell 1993:7).

A Sarracenia purpurea petalis roseis, scapis brevioribus (16.3—35.1 cm longis), floribus

grandioribus (bracchiis stylorum 2.6—4.1 cm longis), labiis ascidiorum crassioribus (2.6-
mm crassis), labiis ascidiorum profunde curvis bases versus ascidiorum in speciminibus
exsiccatis dorsiventraliter complanatis differt.

Perennial herb. Rhizomes usually horizontal, occasionally vertical, 1.2--
5.8 cm long, 0.8—1.8 cm wide. Pitchers decumbent to ascending, in basal
rosettes, 4—9 in each rosette, 5.9—28.1 cm long, 2.3—6.8 cm wide at tube,
2.1—5.3 times as long as wide, hooded terminally, lipped at mouth, tube
arcuate, winged exteriorly on adaxial surface of tube, petiolate. Hood erect,
reniform to broadly cordate, apex usually slightly recurved and thus often
emarginate in dorsiventrally pressed pitchers, otherwise entire, usually coarsely
wavy in abaxial-adaxial plane in life, unconstricted at base and broadly at-

tached to tubular portion of pitcher, 1.8—6.1 cm high, 3.3-13.4 cm wide,
1.5S—3.7 times as wide as high, extending laterally from pitcher mouth 0.8--
4.2 cm on each side, extending basally from pitcher mouth 0.7—2.8 cm on
each side; adaxial surface pubescent with moderately dense, coarse, stiff, decurved
hairs |.0—2.1 mm long, coarsely reticulately veined with veins usually deep
red or purple-red and contrasting with background, background pale green
or pale green suffused with ees ene surface pubescent with moderately
dense, fine, soft, appressed hairs 0.3—0.8 mm long, color as on adaxial sur-

jaceet

face except color of veins usually contrasting less with background color.
Mouth 1|.9—5.3 cm wide; lip 2.6-7.5 mm thick, curved basally in dorsiventrally
pressed pitchers, with maximum distance from horizontal at junction of
lip and wing, this maximum distance 2.4—9.7 mm. Tube obovate in dor-
siventrally pressed pitchers, slightly constricted apically to wide mouth,
gradually tapered basally to narrow petiole, 3.3—17.0 cm long, 2.3—6.8 cm
wide; ventricose in laterally pressed pitchers, strongly outcurved abaxially,
plane or slightly incurved adaxially; pubescent with moderately dense, fine,
soft, appressed hairs 0.3—0.8 mm long except glabrous or sparsely pubes-
cent on abaxial surface; pale green or pale green suffused with dull purple-
red, frequently with longitudinal veins and some cross-veins dull red and
contrasting with ‘cs background. Wing semi-oblanceolate to semicircu-
lar, 3.2—-16.6 cm long, 0.6—5.4 cm wide, pubescence and color as on tube.
Petiole solid, 0.2—6.3 cm long, decurved or straight, glabrous. Flowers
pendent, borne singly on scapes, bracteate. Scape 16.3—35.1 cm high, 1.1—

Naczi eT AL., A new Sarracenia from the southeast 1189

2.8 times as long as longest pitcher per plant, 2.5—-5.4 mm wide at mid-
height, distally abruptly decurved to point of attachment with flower. Bracts
appressed to calyx, 3, broadly ovate, obtuse, entire, 5-8 mm long, 4—7 mm
wide, glabrous, persistent through death of scape and shedding of seeds.
Calyx 5.7—10.6 cm wide in pressed flowers; sepals proximally imbricate,
distally non-overlapping, 5, ovate, obtuse, entire, 3.1—4.7 cm long, 1.7-
3.8 cm wide, 1.4—2.3 times as long as wide, glabrous; adaxial surface usu-
ally very pale green, with deep purple-red margin at anthesis and for a short
time afterward, purple-red margin fading to pale green with age; abaxial
surface usually deep purple-red to pale purple-red, rarely pale green; per-
sistent as long as scape and pericarp remain green. Corolla 8.7—13.2 cm
wide in pressed flowers; petals non-overlapping, 5, pene obtuse, en-
tire, 4.5—6.4 cm long, limb 2.9-4.2 cm long and 2.0—3.5 cm wide, base
1.5—2.3 cm long and 1.5—2.2 cm wide, isthmus Let cm wide, glabrous,
very pale to deep pink or very rarely pale yellow in life, rose to lavender or
rarely nearly white when recently dried, fading to pale brown or whitish
when dry for a longer time, color of adaxial surface same as color of abaxial
surface, shed soon after pollen release. Androecium of numerous stamens,
2.4—3.1 cm wide in pressed flowers; filaments 6-13 mm long; anthers 2.7—
4.0 mm long, 2.0-3.5 mm wide, yellow to red-brown; shed soon after pol-
len release. Gynoecium of 5 connate carpels; ovary globose, 0.9—1.3 cm
high, 1.1—1.4 cm wide, very densely verrucose, pale green or whitish; style
basally styliform, styliform portion 0.8—1.0 cm long, apically abruptly ex-
panded and umbraculiform, persistent through death of scape and shed-
ding of seeds; stigmas 5. Style expansion pentagonal, each angle slightly
elongated and forming a short and emarginate lobe, otherwise entire, 4.8—
7.4m wide, adaxial surface pubescent and glandular-puncticulate, abaxial
surface pubescent and glandular-puncticulate, ribbed, membranous during
anthesis, thickening soon after anthesis and becoming coriaceous, pale green
to nearly white and sometimes lightly suffused with purple-red; lobes 0.4—
1.0 cm long, 0.2—0.3 times as long as style arms, notches 2.9—6.2 mm deep;
hairs on adaxial surface of style expansion moderately dense, erect, fine, 0.2—
0.6 mm long; hairs on abaxial surface of style expansion sparsely to moder-
ately dense, erect, fine, 0.2—-0.5 mm long; glands sparsely to moderately
dense on adaxial surface of style expansion, moderately dense on abaxial surface
of style expansion, slightly raised, depressed-dome-shaped, 0.06—0.13 mm
wide, usually pale to deep red-brown, most prominent during anthesis, becoming
obscure with thickening of style expansion; ribs 5, each oe from center
of style expansion and terminating in stigma; style arms 2.6—4.1 cm long,
0.078—0.20 as long as scapes. Stigmas 0.2—0.6 mm long, 0.4—0.5 mm wide,
each restricted to apex of papilla; pee 5, each in base of notch of style
expansion, perpendicular to lobes of style expansion, 0.8—-1.6 mm long. Capsule

1190 SipA 18(4)
TABLE |. maa characters that best distinguish Swrracenta rosea from S. purpurea. Quantitative
data are means + | SD and elie for measurements. N = number of pager measured. The t

species ditt spear fo all quancit ative har a rs U ,P « 0. Ol). The quantitative rans

ters are listed in descending ae of f-value.

Character S. rosed S. purpurea t-value
1. lip thickness (mm) £54 1.1 1.8 + 0.5 19
(2.6-7.5) (O.7—3.1)
N = 48 N = 85
2. scape height/scyle arm length 8.2 + 1.8 18 + 4.5 18
(4.9-13) (9.5=33)
N = 52 N = 86
3. scape height (cm) 25.9 + 5.03 44.2 + 11.3 13.4
(16.3—35.1) (22.3-79.1)
N N = 86
t. style arm length (cm) 3.2 + 0.34 2.5 + 0.44 9.9
(2.6-4.1) (1.7—3.8)
l= 52 \ = 86
S. lip concavity, maximum 5.0 4 1.7 O.7 + | 8.4
distance from horizontal (mm) (2.4—9.7) (Q—5.3)
N = 34 N = 65
6. hairs on adaxial surface 1.5 4 0.2] 1.2+0.3 7.5
of hood, length (mm) (1.0—2.1) (0.6—2.2)
N = 47 N (oye)
7. petal length (cm) S.1 + 0.48 {1 + 0.45 7.4
(4.5—6.4) (3.3-5.3)
18 25
8. scape height/longese 2.0 + 0.47 3.3.+ 0,74 ic
pitcher length (1.1—2.8) (1.95.2)
N= 17 N = 37
9. sepal length (cm) 3.8 4 0.4] 3,2 + 0.42 6.8
(3.1-4.7) (2.2-4.2)
| = 34 N = 74
10. mouth width (cm) 3.3 4072 2.4 + 0.44 6.5
(1.9-5.3) (1.4—3.6)
N = l i
11. petal width (cm) 2.7 + 0.35 2.1 + 0.32 5.8
(2.0—3.5) (1.6—2.9)
18 N = 25
12. petal color usually pink usually maroon or red
depressed-globose, |.1—1.4 cm high, 1.6—2.1 cm wide, densely verrucose,

brown, dehiscing basipetally. Seeds ae or oblong in outline, compressed,
narrowly ridged along one lateral margin, pyriform in cross-section, 1.8—
2.3 mm long, 1.0-1.4 mm wide, brown, verruculose, often slightly glau-
cous because of a thin coating of wax.

Naczi ET AL., A new Sarracenia from the southeast 1191

Fic. 3. (Left) Sarracenia rosea flowering in the field. Florida, Liberty Co., 15 March 1994,
Naczi 3651.
Fic. 4. (Right) Sarracenia purpurea flowering in the field. Michigan, Chippewa Co., 1995.

Diagnostic Features
Several morphologic features distinguish S. rosea from S. purpurea (Table 1).
The most obvious diagnostic feature is petal color. Sarracenia rosea usually
has pink petals (Fig. 3), whereas 8. purpurea has maroon or red petals (Fig
4). Closely correlated with the paler petals of S. rosea are paler gynoecia (Schnell
1993), though we have observed S. purpurea with pale gynoecia, too. While
Schnell (1993) emphasized the pink coloration of the petals in his diagnosis
of S. purpurea var. burkii, previous researchers believed pink petals were the
result of a mutation (Wherry 1933) or of a phenotypic response to low light
levels (Bell 1949). Through field observations and greenhouse cultivation,
we confirm that pink is the predominant color for petals of 8. rosea. Indeed,
among the species (not hybrids) of Sarracenia, pink petals are unique to S. rased.
In S. rosea, petal color ranges from very pale pink (almost white) to deep
ink. Most commonly, the pink is of medium saturation (as in Fig. 3). Bell
(1949: 157) mentioned observing S. rosea (as S. purpurea) near Bay Minette,
Alabama that had dark red petals. We agree with Schnell (1993) that such
plants are likely introgressants with one of the species that possesses ma-
roon petals, most likely S. /ewcophylla Raf. Putative hybrids of S. /ewcophylla
and S. rosea (as S. purpurea) are reported from several localities, including
Bay Minette (Bell 1949, 1952; Bell & Case 1956; McDaniel 1971). Very
rarely, plants of both S. purpurea and S. rosea produce yellow petals, due to
failure of anthocyanin production (Sheridan & Mills 1998a, 1998b; Hanrahan
& Miller 1998). These yellow-flowered plants also lack reddish coloration

—

1192 Sipa 1&(4)

Fic. 5. Petals of Sarracenia purpurea (top row) and S. rosea (bottom row). Petals pressed and
dried from wild, sun-grown plants. Top row (left to right): Pennsylvania, Bradford Co.,
Naczi 4322; Ontario, Thunder Bay Distr., Garton 18798, New Jersey, Burlington Co., Naczz
156A, 5 Jun 1982. Bottom row (left to right): Florida, Liberty Co., Kra/ 63291; Florida,
Liberty Co., Naczi 3651; Alabama, Mobile Co., Kra/ 16530. Scale bar = Scm.

in pitchers and calyces. This variation of S. purpurea is S. purpurea £. heterophylla
(Eaton) Fernald. For S. rosea, the analogous form has been described by Hanrahan
and Miller (1998), but must be transferred from S. purpurea, as we do here.
Sarracenia rosea f. luteola (Hanrahan & Miller) Naczi, EW.Case, & R.B.Case,
comb. nov. BasionyM: Sarracenia purpurea ssp. venosa var. burkii f. luteola Hanrahan

& Miller, Carniv. Pl. Newsletc. 27:16. 1998
In addition co petal color, we found that nearly every feature of the flower
is substantially larger in S. rosea than in S. purpurea. The sepals of S. rosea are
3.1-4.7 cm long, whereas those of 8. purpurea are only 2.2—4.2 cm long.
The petals of S. rosea are longer and wider than those of S. purpurea (Fig. 5):
4.5—6.4 cm long and 2.0—3.5 cm wide for S. rosea versus 3.3—-5.3 cm long
and 1.6—2.9 cm wide for S. purpurea. Style size, as measured by style arm
length, is greater for S. rosea (2.6-4.1 cm) than S. purpurea [1.7—2.9(—3.8) cm}.
The scapes of S. rosea are relatively short, only 16.3—35.1 cm high versus
22.3-79.1 cm high for S. purpurea. These short scapes are remarkable in

Naczl ET AL., A new Sarracenia from the southeast 1193

Fic. 6. Fruiting plant of Sarracenia purpurea (left) and of holotype of S. rosea (right). Col-

lected from sunny habitats in the wild and pressed and dried. Left: Ontario, Dufferin Co.,
Naczi 4514. Right: Florida, Liberty Co., Naczi 3016. Scale bar = 5cm.

light of the comparatively large flowers of S. rosea. As a result, the ratio of
scape height/style arm length is a particularly strong quantitative character
in separating S. rosea from S. purpurea (Table 1). The values of this ratio are
4.913 for S. rosea and 9.5—33 for S. purpurea. The visual difference between
the two species in flower size-scape height proportions is striking (Fig. 6).
Several vegetative characters are also diagnostic. First, the lips of S. rosea
are thicker than those of S. purpurea (Fig. 7): 2.6-7.5 mm thick for S. rosea
versus 0.7—3.1 mm thick for S. purpurea. Second, in dorsiventrally pressed
pitchers, the lips of S. rosea curve basally, forming a relatively deep concav-
ity. The lips of S. purpurea are horizontal or form a shallow concavity. Thus,
the maximum distance of the lip from horizontal (the maximum depth of
the concavity) is greater in S. rosea than S. purpurea (Fig. 7): 2.4-9.7 mm for
S. rosea versus 0O—5.3 mm for S. purpurea. A third diagnostic feature from
the pitchers is the length of the stiff, decurved hairs on the adaxial surface
of the hoods. In S. rosea, these hairs are (1.0—)1.3—2.1 mm long, whereas
they are 0.6—1.6(—2.2) mm long in S. purpurea. Fourth, the pitchers of S.
rosea are relatively thin-walled, while those of S. purpurea are thicker. Prob-
ably as a result of this difference in thickness, overwintered pitchers of S.
rosea exhibit slight to extensive winter-kill. Pitchers of S. purpurea are ever-
green, even on plants in the northern portion of its range.
The pitchers of S. rosea tend to be larger than those of S. purpurea (Fig. 7).

1194 Sipa (8(4)

Pitchers

SS

Fic. 7. (Left) Pitchers of Sarracenia purpurea (top row) and S. rosea (bottom row
pressed and dried from wild, mature, and sun-grown plants. ae row (left to right): Ontario,
Dufferin Co., Nacz7 4514; New Jersey, Burlington Co., Naez/ 4433; North Carolina, Montgomery
Co., e LOPS: se row (left to right): Florida, Liberty C ‘0., Naczi 30106; Mississippi,
George Co., Naczi 3057; Alabama, Mobile Co., Nacz/ 30606. Pale bar = Sem.

Fic. 8. (Right) ain of Sarracenia rosed. ee pressed and dried from wild, macure,
and sun-grown plants. Top row (left co right): Alabama, Mobile Co., Nacz7 3053; Alabama,
Mobile Co., Nacz7 2809; Florida, Escambia Co., Nacz7 3051. Bottom row (left to right):
Alabama, Mobile Co., Nacz7 3066; Mississippi, George Co., Naczi 3057. Scale bar = Scm.

As a result, the ratio of scape height/longest pitcher length differs for the
two species: 1.1—2.8 for S. rosea and 1.9—5.2 for S. purpurea. Also, pitchers
of S. rosea have wider mouths (1.9-5.3 cm wide) than those of S. purpurea
(1.4—3.6 cm wide). However, we find most pitcher dimensions and shapes
to be practically useless in unambiguously separating S. rosea and S. purpurea.
Pitchers of both species are quite variable in size and shape (Figs. 7, 8).
Measurements of many pitcher characters we studied (pitcher length, pitcher
width, hood height, hood width, width of the portion of the hood that ex-
tends laterally from the mouth, wing width, pitcher length/pitcher width,
hood width/hood height, hood height/pitcher length, hood width/pitcher
length, hood height/mouth width, hood width/mouth wide. mouth width/
pitcher length, mouth width/pitcher width, wing width/pitcher length) overlap
extensively for S. rosea and S. purpurea. The t-values for measurements of
these characters are all lower than for any of the characters listed in Table 1.
Some of the quantitative characters exhibit clinal variation across the
combination of S. purpurea and S. rosea (Fig. 9A). For example, style arm
length is highly correlated with latitude in the combined data set (r =

Naczi eT AL., A new Sarracenia from the southeast 1195

5 T T T 80 T ie) T T
~_—-_-~
z val 0
— . ‘= @)

4b . 4 oO
<= ae) — 6 0F Oo oOo 4
D : fe) fo) ° .

- O

iS ch eel ~ Be°% |
o = 8 oy Oo

ST TF, G00 1 § 80 oP F&
E PE f. ¢ £ ao} a ne
o Po eaaateek oy: @ TS 680°8

fo) fe) OQ 3 0+ O fe) re a

2+ e) 7 oO fe)
= oy 3) ° O
~ M20 © 4
ahoet | he
/7)

1 Il 1 l 10 l l 1

20 30 40 50 60 20 30 4 50 60
A latitude B latitude

Fic. 9. Scatter plots of some diagnostic features vs. latitude for Sarracenia ale (open
circles) and S. rosea (solid squares). A. Cline in style arm length. B. Absence of cline in

scape length.

ce

-0.76, p < 0.001, N = 138), as is length of hairs on the adaxial surface of the
hoods (r = -0.75, p < 0.001, N = 130). However, several other characters do
not exhibit clinal variation (Fig. 9B). For example, scape length has a very
low correlation with latitude (r = 0.377, p < 0.001, N = 138), as does the
ratio of scape length/longest pitcher length (r = 0.34, p = 0.012, N = 54).
The diagnostic characters that are nonclinal indicate S. rosea is not merely
the extreme of a cline of S. purpurea, but rather a taxon distinct from it.

In summary, several characters can be used to separate S. rosea from S,
purpurea (Table 1). The quantitative features that are the best for distin-
guishing the species, based on having the highest ¢-values, are both vegeta-
tive (lip thickness) and reproductive (scape height/style arm length). Though
ranges of measurements for all of the quantitative characters overlap for both
species, a scatter plot of scape height/style arm length versus lip thickness
separates specimens of S. rosea and S. purpurea (Fig. 10). This plot and the
wealth of other diagnostic features for S. rosea, including qualitative char-
acters, Clearly indicate S. rosea is distinct from S. purpurea at the rank of species.

Evidence from additional sources also indicates S. rosea deserves recogni-
tion as a species. In greenhouse cultivation, the differences between S. rosea
and S. purpurea are maintained. Under common greenhouse conditions for
15 years or more, plants from 7 populations of 8. rosea continue to possess
pink petals, short scapes, thick lips, and deeply concave lips while plants of
S. purpurea from 8 populations continue to possess maroon petals, tall scapes,
thin lips, and horizontal or shallowly concave lips, as examples of some of
the differences that are maintained during common cultivation. These re-

1196 Stipa 18(4)

purpurea

length

scape height/style arm

lip thickness ‘

Fic. LO. Scatter _ of scape height/style arm length vs. lip thickness for Sarracenia purpur
and : rosea. Open circles represent specimens of S, purpurea. The solid circle represents the
lectotype of S. aneen plate 70 of Catesby (1738). Solid squares represent specimens of 5.
rosea, with the boxed square depicting the holotype.

sults suggest the differences we have noted between the species are geneti-
cally-based. A second additional source of support for recognizing S. rosect
comes from allozyme analysis of the S. purpurea complex. Godt and Hamrick
(1999) discovered that S. rosea (as S. purpurea var. burkii) is the most geneti-

cally divergent member of the complex. In addition, the genetic distance
between S. rosea and S. purpurea is relatively large and is similar to that found
between many congeneric species (Gottlieb 1977, 1981; Crawford 1983).
Nomenclature

We are unaware of any prior names at the rank of species that apply to S.
rosea. None of Rafinesque’s (1840) names for Sarracenia (as “Sarazina’)

Naczi eT AL., A new Sarracenia from the southeast 1197

appear to apply to S. rosea, though the applications of these names are some
what uncertain. Geographically, four of these new names could fit S. rosea.
Rafinesque stated his S. venosa, “...differs from S. gébbosa {a northern-rang-
ing new species of his, under which he lists S. purpurea as a synonym], by ...
flowers smaller—Virg. ad Florida.” Clearly, S. venosa cannot apply to the
large-flowered S. rosea. As well, S. parviflora Raf. (“... appendice [hood} concavo
fornicato ... very distinct sp. of Florida, yet akin to the last [§. venosa}, leaves
3 to 6 inches long, nervose, flowers very small, purplish.”) does not apply
to S. rosea. Most likely, S. parviflora is a synonym of S. psittacina Michx. (a
quite distinctive species having small flowers and relatively short pitchers
with strongly arched and concave hoods), though McDaniel (1971) placed
S. parviflora in synonymy with S. purpurea. Both S. acuta Raf. (“fol. tubul.
longiss. nervosis ala angustissima ... Alabama ...”) and S. adunca Raf. (“... fol.
tubul. longis ... ala angust. ... Florida ...”) do not apply because S. rosea
has short pitchers with wide wings. Also, 8. adunca Raf. is preoccupied by
S. adunca Sm., 1805, a synonym of S. minor Walter. Unfortunately, bota-
nists probably will never know the application of Rafinesque’s names in
Sarracenia.Most of Rafinesque’s herbarium was discarded soon after his death
in 1840 (Stuckey 1971). Previous workers have not located types of Rafinesque’s
names in Sarracenia. oe by us and herbarium curators for Rafinesque
specimens at DWC, G, NY, P, PH, and WIS have been fruitless.
McDaniel (1971) ae S. purpurea with plate 70 of The Natural
History of Carolina, Florida and the Bahama Islands (Catesby 1738). Catesby
did not mention the provenance of the illustrated specimen, but most likely
it was outside the range of S. rosea. His “Map of Carolina, Florida and the
Bahama Islands, with the Adjacent Parts” indicates the Florida of his day
ranged only as far west as the present-day Aucilla River, east of the range of
S. rosea. Nevertheless, the possibility that Catesby obtained the plant from
further west means the plate must be evaluated before S. rosea can be ac-
cepted as a correct name. Although the petals are anomalously purplish pink
(though described by Catesby as “...of a purple colour...” in text accompa-
nying plate 70), the species illustrated in the plate is otherwise typical S.
purpurea. Measurements of the plate for nearly every diagnostic character
lie within the range for S. purpurea, but several are outside the range for S.
rosea. For example, lip thickness is 1.2 cm, style arm length is 2.1 cm, and
the ratio of scape height/style arm length is 16. In addition, the lips appear
to be horizontal, not concave, though the partially obscuring hoods make
the determination of this condition somewhat uncertain. Measurements of
the Catesby plate fall well within the cluster of specimens of S. purpurea in
the plot of scape height/style arm length versus lip thickness (Fig. 10).

on)

—<

198 Sipa 18(4)

Etymology
We have chosen “rosea,” meaning pink, as the epithet for this new species
because of its distinctively colored petals. This epithet is descriptive, easily
comprehended, and is in the tradition of several other specific epithets in
the genus by applying to flower color (e.g. S. flava L., 8. rubra Walter, and
S. purpurea). We use a new epithet for this species, rather than transfer the
epithet used when this taxon was treated as a variety (Schnell 1993), as permitted
by article 11.2 of the International Code of Botanical Nomenclature (Greuter et
1994). Our choice is intended to avoid confusion of ranks that may en-
sue from using the same epithet for both variety and species. Furthermore,
by using a new epithet, we underscore the recognition of S. rosea as a spe-
cies with a suite of features that differentiate it from S. purpurea.

Phenology

The flowering period of 8. rosea extends from mid-March to mid-April, with
the peak in the last 10 days of March. The earliest flowering date of her-
barium specimens is 14 March and the latest is 20 April. At the time of
Howering, pitchers of the current season are usually undeveloped or incom-
pletely developed. Overwintered pitchers are present at the time of flowering,
but these are often slightly to nearly completely brown and withered due
to winter-kill (Fig. 3)

Typification

For the holotype of 8. rosea (Fig. 6), we have selected a post-anthesis speci-
men because it has fully developed pitchers of the current season. In addi-
tion, the holotype has the thickened style expansion typical of post-flowering
plants. Such a style expansion is less prone to damage than the fragile ones
of plants during anthesis. The holotype is quite representative of S. rosea for
key quantitative features (Fig. 10). Plants observed from the same popula-
tion as the holotype in the following year uniformly bore flowers with pink
petals (Naczz 3639).

Distribution

Sarracenia rosea ranges from southwestern Georgia (Tift County) and north-
western Florida (Gadsden County) west to George and Jackson counties,
southeasternmost Mississippi (Fig. 11). Ic occurs solely on the Coastal Plain
in the drainage of the Gulf of Mexico. Most populations are within 120 km
(75 mi) of the coastline. McDaniel (1966) cites a specimen of S. purpurea
from Taylor County, Georgia, which we have not seen, that is probably S.
rosea. The collection locality lies within the Gulf of Mexico drainage, but it
is farther norch than any other population known and about 300 km (185
mi) from the coast. The range of S. rosea is almost identical to that of S.
lencophylla (McDaniel 1971), with which it often grows.

=

—

Naczi er AL., A new Sarracenia from the southeast 1199

Sarracenia rosea

100 km

Fic. 11. Distribution of Sarracenia rosea, based on voucher specimens we examined.

Sarracenia rosea and S. purpurea are allopatric. Sarracenia purpurea ranges
further north and east of S. rosea. The collection of S. purpurea that is closest
to the range of S. rosea is from Tattnall County, Georgia (Harper 2151). Thus,
a minor disjunction of about 135 km (85 mi) separates the ranges of S. rosea
and S$. purpurea. More significantly, S. purpurea is apparently absent from
the Gulf of Mexico drainage, the Tattnall County site and all other locali-
ties in the southeastern U.S.A. being in the Atlantic drainage.

Based on two specimens, S. purpurea is reported from Louisiana (MacRoberts
& MacRoberts 1988). One would expect these collections to be S. rosea, since
Louisiana is in the Gulf of Mexico drainage and far from the range of S.
purpurea. The attribution of S. reser to Louisiana based on these collections
is problematic, though. The original label of one of these collections (US
782242) reads, “Sarracenia purpurea/wet pine woods/St. Helena, La.” With
its lip 1.4 mm thick, lip’s maximum distance from horizontal of 1.6 mm,
scape height of 42.4 cm, and style arm length of 2.4 cm, this collection is
clearly 8. purpurea, not S. rosea. The second collection reputed to be a voucher
from Louisiana (LSU 006607) has “Saraceniaceae [sic }/Sarracenia rubra Walt./
Red flowered Trumpetleat/Covington/AE [“AL.”?}" handwritten on the original
label. The specimen is quite fragmentary; it consists of two isolated, later-
ally pressed pitchers and a portion of a scape bearing only 2 sepals (3 sepals,
corolla, androecium, and gynoecium are all lacking). This specimen is also

1200 Sipa 18(4)

S. purpurea, since the lip is only 1.1 mm thick. The scape fragment is 30.1
2

cm high and the longer sepal is 3.3 cm long, both of which are inconclu-
sive since these measurements fall within the range of overlap between S.
purpurea and S. rosea. The collection locality of this specimen may not be

Louisiana. The script leaves determination of the final pair of letters equivocal.

Instead of “...Covington/AF. [Americus Featherman, a collector in late 1 9th-
century Louisiana}” (MacRoberts & MacRoberts 1988), the label may ac-
tually read “...Covington/AL.” Thus, the label may actually indicate Covington
County, Alabama. The single period after the final pair of letters (wor after
each letter, as one would expect if the correct reading is “A.E.”) supports
this interpretation. Further support is the fact that S$. rvbra is unknown from
Louisiana (Murry & Urbatsch 1979), but does occur in Covington County,
Alabama (e.g., MacDonald 13556, IBE). Whatever its origin, the specimen
does not appear to belong with its label since it is neither S$. rvbra nor S.
rosea, Which would be expected if the collection came from either Covington,
Louisiana or Covington County, Alabama. In light of the evidence, we ad-
vise caution when considering the inclusion of $. purpurea in the flora of
Louisiana. It seems more reasonable to discount the two specimens because
of possible labeling errors than to accept the disjunct occurrence of 8. purpurea
in Louisiana, a disjunction of about 800 km (S00 mi) southwest from the
nearest population in Tattnall County, Georgia.

Habitats
Sarracenia rosea usually grows in sunny to lightly shaded, wet, sandy, acid

on

soil in relatively open sites, often with scattered pines and shrubs. Popula-
tions of S. rosea also occur in ditches, shrubby thickets, edges of swamps,
and the dense shade of swamp interiors. In swamps, population densities
and the proportion of flowering or fruiting plants are lower than in sunnier
habitats. Using the terminology for pitcher plant habitats of Folkerts (1991),
most populations of 8. rosea we have observed inhabit seepage bogs and sa-
vannas, though we have also observed them in stream terraces and swales.
Usually, S. rosea grows with other Sarracenia species. We have observed it
growing with S. alabamensis BW.Case & R.B.Case ssp. wherryi FE W.Case &
R.B.Case, S. alata (A.W.Wood) A.W. Wood, S. flava, 8. leucophylla, 8. psittacina,
and S$. rubra ssp. gulfensis D.E.Schnell.

Conservation

Most populations of S. rosea are in the western panhandle of Florida and
adjacent southernmost Alabama. It is very rare in Mississippi. We have seen
neither populations nor recent collections from Georgia. Where they oc-

cur, plants of S. rosea can be numerous. However, populations are infrequent
and local in the small geographic range of 8. rosea.

Naczl ET AL., A new Sarracenia from the southeast 1201

Folkerts (1977, 1982, 1990) has documented the extensive destruction
of pitcher plant habitats and the consequent decline of pitcher plant abun-
dance. Our experiences have shown habitat destruction and fire suppres-
sion to be among the most serious threats to S. rosea. Many other menaces
exist, though. Despite the avai ety of reasonably priced, commercially
cultivated plants of S. rosea, overcollection from the wild damages many
populations. Twice, when visiting a locality where we had studied S. rosea
populations, we discovered recently-dug holes and most of the plants miss-
ing. Since S. rosea appears to be an uncommon species, 1s exceptionally at-
tractive, and faces many threats to its survival, we recommend its current
conservation status be reviewed and it be considered for protection throughout

its range.
KEY TO SARRACENIA ROSEA AND PURPUREA

Petals usually pink, lip 2.6-7.5 mm thick at thickest point, lip usually ree
curved toward ie base, scape 16.3—35.1 cm high, style arm 2.6-4.1 cn

long, scape height/style arm length = 4.9-13 ee rosea
Petals usually maroon or red, lip 0.7—3.1 mm thick at thickest point, lip usu-
ally horizontal ors palo curved toward pitcher base, scape 22-79 cm high,
style arm 1.7—2.9(—3.8) cm long, scape height/style arm length = 9.5—3

Sarracenia purpurea

Representative Specimens
Specimens marked with an asterisk are those we measured for statistical analyses. U.S.A.
ALABAMA. Baldwin Co.: Gateswood, Passe ne tay sm. (NY, US*); near Bay Minet
30 May 1932, Thuts.n. (OS); 2 mi W | route 90, 3 Apr 1966, Kral 261 i
(IBE, MICH*, VDB*); 5 mi N - Romar Beach, 26 jul 1969, Ellis 90 (VDB*); ca. 5 mi S
of Foley on route 56, 19 Apr 1970, Kral 38766 (VDB*); 2.6 mi S of Bay Minette, 18 May
1975, LeLong 8482 (USAM); Gulf shores, 22 May 1975, Kra/ 55708 (VDB*); Along high-
way 90, between Elsanor and Seminole, 23 Aug 1975, LeLong 8631 (USAM); Off S side
route 90, 14 May 1983, Wilhelm 11333 (VDB*); N of Perdido, 18 Aug nae Naczi 691 &
Folkerts (RANK); By route 112, ca. 10 mi N of Perdido River bridge, 3 May 1987, Kral
73905 (VDB*); ca. 11 mi N of Stockton, 17 May 1987, Naczi 1605 (KNK); N of Elberta,
28 ae 1988, Naczi 1803 (KNK); Perdido, 22 May 1993, Naczi 3070 (KNK*). Coffee

1.5 mi NE of Kinston, 9 i 1932, Wherry s.n. (PH). Conecuh Co.: Highway 31 at
Travis i Sepulga River, 2 Aug 1985, Diamond 1367 (AUA), mann Co.: high-
way 4, 13 1998, Mac ce 11048 (BE). Escambia Co.: ca. 1.5 mi E of Wawbeek, :

Jun 1969, aoe 147(1BE, VDB*); interstate 65 between routes 21 ae = 7,16 May 1971,
Evans 46180 (NCU, TENN, VDB*); Little River State Park, 30 May 1982, Whilhelm 10132
(VDB*); WNW of Flomaton, S of Big Escambia Creek, 27 Jun 1993, Anderson 14411 (FSU).
Geneva Co.: 2 mi S of Hartford, 14 May 1972, Hanning155 (AUA); ca. 9 mi SW of Samson,
29 May 1998, MacDonald 11314 (BE). Mobile Co.: Whistler, 18 Apr 1895, Modr s.n.
(US 782237*); Whistler, 18 Apr 1895, Modr s.n. (US 782240*); Whistler, Mar 1918, Graves
sn. (MO*); Irvington, 30 Mar 1951, Liggett 1711 (FLAS*); ca. 7 mi W of Citronelle, 7
Apr 1963, Kral 16530 (FSU, VDB*); 3.3 mi N of Citronelle, by route 45, 3 Jun 1970,
Kral 39598 (VDB*); Along highway 188 to Coden, 23 Mar 1974, Lelong 7450 (USAM,
VDB*); W of Citronelle, along Escatawpa River, 30 Mar 1974, Le/ong 7512 (USAM, VDB*);

1202 Sipa 18(4)

patos

nae highway 90, E of state line, 24 Mar 1976, Lelong 9046 (USAM*); ESE of Bayou La
Batre, 29 Apr 1988, Naczi 1804 (KNK ), 27 Apr 1991, Naczi 2809 (KNK), 21 May 1993,
ae 3053 (KNK*); W of Citronelle, 22 May 1993, Naczz 3066 (KNK*). FLORIDA.
Bay Co.: Mill Bayou, St. Andrews Bay, 17 Mar 1926, Banker 3556 (NY); Youngstown, 16
Mar 1937, Exploration Party 1937 (FLAS); Panama City, 20 Mar 1943, Sargent s.n. (US*);
8 mi E of Panama City, 25 Mar 1949, Hood 1699 (FLAS); Parker, 15 Apr 1971, Athey 5.7.
Say ee Co.: Myron, 5 May 1930, Moldenke 1158 (MO*, NY); ca. 0.5 mi N of
nard, 14 Sep oe ees 2382 (FLAS*). Escambia Co.: Bluff Springs, 13 Apr 1935,
Th s.m. (FLAS); 3 mi N of Cantonment, 22 Jun 1963, McDaniel 3553 (BE); 3 mi ESE
of Pleasant enn 5 ce 978, Hansen & Hansen 4784 (FSU*); Pensacola, 19 May 1993,
Naczi 3051 (KNK*). Gadsden Co.: inter Quincy et Aspalaga, in pinetis, May 1843, Rigel
sm. (US*), in uliginosis ad rivulos, May 1843, Ruge/ sn. (FLAS, NY); [no additional local -
ity data], [no date], seas n. (NY); SW of Quincy, 23 Apr 1924, Small 11214, DeWinkeler,
& Mosier (NY). Holmes Co.: E of Ponce de Leon, 29 Nov 1929, Hume s.n. (FLAS); Ponce
de Leon, 18 apes 1937, splat ee 1937 (FLAS*); SW of Leonia, 4 Apr 1958, Godfrey
56393 (FSU*, NCU). Liberty Co.: 6 mi N of Vilas, 26 Feb 1956, Kral 1965, Godfrey, &
Kurz (FSU); 4 mi N of Sumatra, 6 Oct 1 — se eee (IBE); 4.2 mi N of Wilma, 6
May pe es 57908 aon VDB*); Kra/ 63291 (VDB); S of junction of route 65 and

route 20, 20 May 1976, Solomon 2159 (MO*); “ of Wilma, 20 May 1976, Solomon 2180
(MO*); ne 3 mi S of Hosford, 6 Aug | se oe 18 (USCH*); 6 mi S of Telogia, 16
Apr 1987, Godfrey 82347 (FSU*); S of Wil May 1987, Naczi 1614 (KNK); SW of

Telogia [type locality, 1 4 Mar 1994, oe z/ ae ea N of Wilma, 15 Mar 1994,
Naczi 3651 (RNK*, MICH*, MO*, NY*), 4 Aug 1 ae Sie wee (KNK*, peat S
of Wilma, 4 Aug 1994, Naczi 4503 (KNK*). Okaloosa Co.: 4 mi N of Dorcas, oe
1954, Ford 3973 (FLAS); ca. 3 mi SW of Crestview, aa 1960, aan sin. (FLA

mi E of Crestview, 7 Apr 1963, Godfrey 62693 (FSU*); Eglin Air Force Base, Malone ne "
30 Jul 1992, Anderson ey (FSU); ca. 5 mi NW of Niceville, 18 May 1993, Naczi 3026
(KNK*, MICH*); ca. 3.5 mi S$ of Crestview, 18 May 1993, Naczi 3027 (KNK*); Eglin
Air Force Base, Metts Creek, 24 Mar 1994, Anderson 14698 (FSU). Santa Rosa Co.: N of
Holt, 20 Apr nen Godfrey 59425, pee & Henderson FSU, NCU); ca. 11 mi N of Holley,
Eglin Air Force Reservation, 21 Jun 1967, Smith 1549 (FLAS); ca. 6 mi N of Holley, 21
Jul 1977 eae 329 (FLAS); NE of Holley, Eglin Air Force Base, 27 Mar 1994, Anderson
14723 (FSU). Walton Co.: [no additonal locality data}, summer 1885, Curtiss s.n. (NY);
N of DeFuniak Springs, 30 Mar 1949, Hood 1784 (FLAS); ca. 12 mi S of DeFuniak Springs,
4 Mar 1956, Godfrey 54407 (FSU, NY); ca. 14 mi NE of Niceville, Eglin Air Force Reser-
vation, 29 Jun 1966, Beckner 1429 (FLAS*); ca. 2.5 mi S of Mossy Head, Eglin Air Force
Reservation, 23 Oct 1966, Chapman 171 & es (FLAS). Washington Co.: 4-5 mi
SW of Chipley, 16 Apr 1937, Hobbs 5.2. (FLAS). C RGIA. Randolph Co.: SW of Coleman,
28 Oct 1902, Harper 1783 (NY, US*). Tift es aes {now Sunsweet], 4 Feb 1892,
Rolfs (FLAS). MISSISSIPPI. George Co.: S of Agricola, 28 Mar 1964, McDaniel 4044
(IBE); Movella, 21 May 1993, Naezi 3057 (RNK*). Jackson Co.: ca. 1.5 mi NE of Or-
ange Grove, near Jackson Creek, 16 Jun 1963, McDaniel 3539 (IBE, TENN*).

~~

APPENDIX
Measured Specimens of Sarracenia purpurea
CANADA. NEW BRUNSWICK. Restigouche Co.: Charlo, | Aug 1894, Fow/ler s.n. (US). NEW-
FOUNDLAND. Whitbourne, 16 Aug 1894, Robinson & Schrenk 64 (US). 15 mi W of Gander, along
shoreline of Gander River, 16 Jul 1949, Bassett 498 (MO). Placentia West Distr.: Pipers Hole River
3ridge, route 11, 25 Jul 1960, Rowlean 5770 (US). St. Mary’s Distr.: ca. 3.5 mi N of Riverhead, 16
Aug 1961, Rowleaw 7008 (US). Labrador: Goose Bay, 26 Jun 1950, Gillett 5123 & Findlay (US).

Naczl ET AL., A new Sarracenia from the southeast 1203

NOVA SCOTIA. Canso, 12 Jul 1901, Fowler 5.x. (US). Cape Breton National Park, west side, 1S
1947, Swallen 9880 (US). Inverness Co.: Cape Breton Highlands National Park, White Capes re-
gion, 16 Jul 1970, Ustal 7415 (FSU). ONTARIO. Algoma Distr.: 3 mi past Anjigami along road
following pee 20 mi NE of highway 17, T28, R22, 47° 53’N, 84° 37°W, 15 Aug 1971, Garton
14741 et al, (MICH). Bruce Co.: 8 mi N of Southampton, South Sauble Beach, in Silver Lake, 17 Jun
1948, Soper 3935 & Dale (US). Carleton Co.: Carlsbad Springs, 25 Jun 1947, Cody & Calder 517
(US). Dufferin Co.: ca. 5 mi NW of Grand Valley, Lucher Marsh Wildlife Area, Wylde Lake, 19 Aug
1994, Naczi 4514 (SNK). Parry Sound Distr.: Opposite sakes Township, Big Island, off Eagle
Lake, 23 Jul ee a. 244 (US). Renfrew Co.: 5 mi E of Deux Rivieres, Gibson Lake, along
S side of route 17, n 1987, Naczi 1701 (KNK). Thunder Bay Distr.: Sibley Provincial Park, SE
side of § yes Se back of Middlebrun Bay, 20 Jul 1961, Voss 10138 (MICH); ca. 12-14 mi
nitouwadge, 10 mi N of Nama Creek bridge on Manitouwadge-Caramat Industrial Road,
19 Jul a Voss 11639 (MICH); Thunder Bay City, just N of Northwood Park area, 26 Jun 1975,
Garton 16525 (MICH); 18 km. NW of Dorion on Spruce River cut-off, 1 Jul 1979, Garton 18798
(MICH). Re ee 2.1 mi S of junction of Watson Road and Arkell Road in Arkell, 18 aC
1994, Naczi 4513 (KNK). QUEBEC. Magdalen Islands, Grindstone Island, Grindstone, 17 Jul 1s
Fernald 7531 et al. ao SAINT-PIERRE ET MIQUELON. Ile Saint-Pierre, vicinité de ville rie
Pierre, 24 Nov 1993, Etcheberry s.n. (KNK). U.S.A. CONNECTICUT. Tolland Co.: Stafford, Jul
1898, Morris s.n. (US). DELAWARE. Sussex Co.: E of ek ae 21 Jun 1924, dan 12115
(US). GEORGIA. Tattnall Co.: Sand-hills of Ohoopee River near Reidsville, 26 Apr 1904, Harper
2151 (MO). INDIANA. Wells Co.: Jackson Township, 24 Jul 7605, Deam 91 (US). MAINE. Hancock
Co.: Between Brooklin and Sedgwick, 8 Jul 1915, Safford 83 (US). MASSACHUSETTS. Norfolk
Co.: Canton, 26 May 1894, aaa: sm. (MQ); Canton, 11 Aug 1926, Standley & Blake s.n. (US).
MICHIGAN. Cheboygan Co.: N of Riggsville, 8 Jul 1986, ek zi 1400 (KNK). Livingston Co
ca. 4.5 mi W of Brighton, 22 Aug 1994, nee 4515 (KNK). Mackinac Co.: ca. 5 mi N of Epoufette,
6 Jul 1987, Naczi 1742 (RNK). ae OTA. Cass Co.: Gull ae Jun 1893, ee (US).
St. Louis Co.: 18 mi N of Duluth, 18 a fe ieee 2983 (MO). NEW HAMPSHIRE. Coés
Co.: Shelburne, — 1915, Dest n. hres eshire Co.: Chesterfield, 21 Jul 1972, Soap
(MO). NEW JERSEY. Atlantic Co.: ie ant Mills, 26 May 1932, Hermann 3054 (MICH),
Pleasant Mills, - Nov 1993, Nova & Naczi (KNK). Bergen Co.: Demarest, 30 May 1876,
Brown 1294 (MICH). Burlington Co.: Martha, along E side of Oswego River, 5 Jun 1982, Naczz
156A (KNK), 26 Nov 1993, Naczi 3572 & Naczi (KNK); ca. 4 mi N of Chatsworth, 20 Nov 1993,
Naczi 3563 & Naczi (KNK); Quaker Bridge, ca. 5 mi N of Batsto, 21 Jun 1994, Naczi 4432 & Thieret
(KNK), Naczi 4433 & Thieret (RNK), Naczi 443 4 & Thieret (KNK). Camden Co.: Spring Garden, 4
Jul 1868, Redfield s.n. (MO). Cumberland Co.: ca. 2 mi N of Newport, 21 Jun 1994, Naczi 4392 &
Thieret (KNK). Gloucester Co.: Malaga, 15 Aug 1 oe Adams 458 (MO). Middlesex Co.: 2 km N of
Helmetta, 6 Jun 1966, Kane & Coyne s.n. (US). Morris Co.: Succasunna Jn 1909, Mackenzie 4119

—~

Es
Bi

(US). Och ee Pans 30 pas noe Dowell 3670 (MO), ca. 4 mi S of Whiting, 26 Nov 1993,
Naczi 3 ). Sussex Co.: Budd’s Lake, 12-14 Aug 1890, a n. (US). N EW YOR K.
Onon daga Co.: Cicero ak Jun 1901, House s.n. (US). Soe eenel . Taborton, Cranberry Vly

Creek, 25 Jul 1994, Tucker 10058 & Miller (KNK). NORTH CAR ee Brunswick Co.:
of Bre yea Columbus Co.: 7 May 1965, Chen 442 (USCH); ca. . mi NW ne 20 Dec 1994,
Naczi 4539 (KNK); 5.0 mi NW of Supply, 20 Dec 1994, Naczi 4541 (SNK). Catawba Co.: Near
Hickory, 23 1893, Heller 1014 (MO). Carteret Co.: near the town of Ser 24 Aug 1979
thal s.n. (USCH). Columbus Co.: Delco, 26 Jul 1938, Br s.m. (US). Harnett Co.: Lillington,

24 Mar 1966, Coyne io n. (US). Henderson Co.: swamps of Muddy Creek, 21 Aug 1881, Smith
SM. . ca. 10.5 mi SW of oe 12 Dec 1994, cone G Hill sei (KNK). Jackson Co.:
S of Cashiers, 11 Dec 1994, Naczi 4519 et al. (RNK). Montgomery Co. ack Ankle, 26 May 1988,

Naezi 1975 (KNK). New Hanover Co.: near Wilmington, Apr 1888, ie s.m. (US); S of Kirkland,
15 May 1930, Moldenke 12 238 (US). Onslow Co.: 3.5 mi SW of Maysville, 28 May 1977, Kral 60229
(VDB). Pender Co.: 1 mi N of Burgaw, 11 May 1960, Kra/ 10101 (VDB),; Holly Shelter Wildlife
Refuge, 19 May 1 962, Kral 14723 (VDB); Burgaw bog, 6 May 1968, Wiges ae ia SM. ae
Robeson Co.: N side of Lumberton off route 74, 27 Aug 1963, Kral 19135 (VDB). Tyrell Co.

Kilkenny, 8 May 1938, Kerr & Godfrey 3864 (US). OHIO. Lorain Co.: Camden, 18 Jun 1894, Oberlin
sn. (US). PENNSYLVANIA. Bradford Co.: ca. 2.2 mi SSW of Leroy, Barclay Mountain, 17 Jun

1204 Sipa 18(4)
1994, Naczi 4322 & Thieret IKNK). Tioga Co.: ca. 3 mi S of Leetonia, 25 Jun 1994, Keener 94026
(KNK). SOUTH CAROLINA. Chesterfield Co.: SSW of Cheraw, 20 Dec 1994, Naczi 4523
(SNS). Darlington Co.: Hartsville, ei i 1920, he SM eo Hartsville, Along E side of Kilgore’s
Branch, 20 Jul L941, Smith 877 (USCH). ae ae own Co.: 5.5 mi $ of Georgetown, 29 Jun 1939

Godfrey & Tryon 211 (MO). aie Co.: E of Caesars He " 11 Dec 1994, Naczi 4518 et al. (RNK),
E of Caesars Head, 31 May 1995, Naczi 4755A & Naczi (RNR). Horry Co.: Cotton Patch Bay area,
19 May 1987, Batson & ba sa. (USC . Kershaw Co.: E of Camden, 25 Jun 1984, Sa/im et al.
4 (USCH). Lexington Co.: Platte Spring's Road, 4 May 39, Hechenbleikner s.n. (USCH). VIRGINIA.
Chesterfield Co.: near Chester, 6 Jun 1936, Wilmouth (US). Greensville Co.: ca. | mi NW of Dahlia,
7 Jun 1946, Fernald & Moore ee (MO).

ACKNOWLEDGMENTS

We are very grateful to Bruce A. Ford and Anton A. Reznicek for their
many insightful comments on the manuscript. We thank George W. Folkerts,
T. Lawrence Mellichamp, Edward G. Voss, Anton A. Reznicek, Ken L. Gordon,
Donald E. Schnell, Robert K, Godfrey, Richard M. Eilers, and Charles N.
Horn for sharing locations of populations of 8. purpurea and S. rosea. Rowell
Bosse, Richard M. Eilers, George W. Folkerts, James H. Hill, Charles E.
Moore, Mary C. and Robert F. Naczi, Mary A. Naczi, and John W. Thieret
assisted with field work. Louise Kirn, Ray Hart, and Andrew Colaninno
processed a permit for our collections from Apalachicola National Forest.
Roger Etcheberry, Carl S$. Keener, and Gordon C. Tucker generously col-
lected S. purpurea specimens and sent them to us for our study. We also thank
the curators of AUA, DHL, FLAS, FSU, IBE, KNK, KY, LSU, MICH, MO,
NCU, NY, OS, PH, TENN, US, USAM, USCH, VDB, and WKU for loans
of specimens or assistance during our visits. We particularly thank John R,
MacDonald (IBE) for sharing label data from his collections, and John MacDonald
and Cynthia M. Morton (AUA) for special assistance with specimens. We
appreciate the help of curators who searched for Rafinesque specimens for
us: S. Barrier (P), Theodore S. Cochrane (WIS), Fernand Jacquemoud (G),
P. Morat (P), Martha A. Potvin (DWC), and Rodolphe Spichiger (G). Sheila
Bowling and Brenda Racke measured some specimens. Financial support
for some of the field work came from Northern Kentucky University and
the University of Michigan Rackham School of Graduate Studies.

REFERENCES
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tchell Sci. Soc. 65:137-166.

= c -R. 1952 a brids in the genus Sarracenia: 1. History, distribution, and tax-

eres : Elisha Mitchell Sci. Soc. 68: 55-80.
BELL nd FW. Case. 1956. Natural hybrids in the genus Sarracenia: I. Current notes

on eee Je tie Mitchell Sci. Soc. Peto 52.

-ASE, F.W. and R.B. Case. 1976. The Sarracenia rubra complex. Rhodora 78:270—325.
am ESBY, M. 7 738. The natural history of Carolina, Florida and the Bahama Islands, vol. 2

Printed at the expense of the author, London.

Naczi ET AL., A new Sarracenia from the southeast 1205

Copy, W.J. and $.S. Tatsor. 1973. The pitcher plant Sarracenia alas L. in the north-
western part of its range. Canad. Field-Naturalist 87:31

Crawrorb, D.J. 1983. Phylogenetic and systematic oe a electrophoretic stud-
ies. In: Tanksley, S.D. and T.J. Orton, eds. Isozymes in plant genetics and breeding, Part
A. Elsevier, Amsterdam. Pp. 257-287.

Fotkerts, G.W. 1977. Endangered and threatened carnivorous plants of North America.
In: Prance, G.T. and T.S. Elias, eds. Extinction is Forever. New York Botanical Garden,
Bronx. Pp. 301-313.

Foikerts, G.W. 1982. The Gulf Coast pitcher plant bogs. Amer. Sci. 70:260—267.

Fotkerts, G.W. 1990. The white-copped pitcher plant
Oryx 24:201-207.

Foikerts, G.W. 1991. A preliminary classification of plant habitats in the south-
eastern United States. J. Alabama Acad. Sci. 62:199-225.

a case of precarous abundance.

Gopt, M.J.W. and J.L. Hamrick. 1999. one tic divergence among infraspecific taxa of
Sarracenia pops Syst. Bot. 23:427-438.
Gort Electrophoretic evidence and plant systematics. Ann. Missouri Bot.

ea eee ;
Gortuiss, L.D. 1981. Electrophoretic evidence and plant populations. Progr. Phycochem
7:1-46.
Greuter, W., FR. Barrie, H.M. Burper, W.G. CHALONER, V. DEMOULIN, D.L. HawKswortH,
.M. JORGENSEN, D.H. Nicoison, P.C. Sitva, P. TREHANE, and J. McNem. 1994. Inter-
national code of botanical nomenclature (Tokyo Code). oa ee 1 9

et alas B. and J. Miter. 1998. History of discovery: yellow-flowered Sarracenia purpurea
L. subsp. venosa (Raf.) Wherry var. burkii. Carniv. Pl. Newslett. 27:14-17.
HOLMGREN, ., N.H. Hoitmcren, and L.C. Barnett (editors). 1990. Index Herbariorum.

Part I. The Ferbatia of the World, 8th ed. New York Botanical Garden, Bronx.

Lioyb, FE. 1976. The carnivorous plants. Dover Publications, New York

Macrar ane, J.M. 1908. Sarraceniaceae. In: Engler, A., ed. Das Pflanzenreich, IV. 110, 1—
39. Wilhelm Engelmann, Leipzig.

MacRoberts, M.H. and B.R. MacRoserts. 1988. A note on Sarracenia purpurea L. in Loui-
siana, bas 65:191-194.

McDan S. 1966. A taxonomic revision of Sarracenia (Sarraceniaceae). Ph.D. disserta-
tion, oe State Univ.

McDanikL, S. 1971. The genus Sarracenia (Sarraceniaceae). Bull. Tall Timbers Res. Sta. 9:1—
36

36.

Murry, R.E. and L.E. Ursarscu. 1979. Preliminary reports on ene flora of Louisiana. HI.
The families Droseraceae and Sarraceniaceae. Castanea 44:24—27

RAFINESQUE, C.S. 1840. Autikon ee Philadelphia.

Rou eau, E. and G. Lamoureux. 1992. Atlas of the vascular plants of the Island of New-
(oundland and of the Islands ofS Mie acion. Fleubec, Saint-Henri-de-Lévis,
Québec.

SCHNELL, D.E. 1979. A critical review of published variants of Sarracenia purpurea L. Casta-
nea 44:47-59.

SCHNELL, D.E. 1993. Sarracenia purpurea L. ssp. venosa (Raf.) Wherry var. burkii Schnell
(Saneacenines ety of the Gulf Coastal Plain. Rhodora 95:6-10.

SCHNELL, D.E. and R.O. ee 1997. Sarracenia purpurea L. ssp. venosa (Raf.) Wherry
var. montana Sie & Determann (Sarraceniaceae): a new variety. Castanea 62:60-62.

SHERIDAN, P. M. and R.R. Mitis. 1998a. Genetics of anthocyanin deficiency in Sarracenia
L. HortScience 33:1042—1045.

1206 Sipa 18(4)
SHERIDAN, P.M. and R.R. Mus. 1998b. Presence of proancthocyanidins in mutant green
Sarracenta indicate blockage in lace anthocyanin biosynthesis between leucocyanidin and
pseudobase. Pl. Science 135:11-16.

Sruckey, R.L. 1971. The firse public auction of an American herbarium ee an ac-
count : aie fate ko the Baldwin, Collins, and Rafinesque herbaria. Taxon 20:443—459.
WHERRY, E.T. 193:

3. The geographic relations of Sarracenia purpurea. Bartonia 15: ae
WILKINSON, L

me SYSTAT: the system of statistics. SYSTAT, Evantson, IL.

THE VASCULAR FLORA OF BISCAYNE
NATIONAL PARK, FLORIDA

RICHARD STALTER and JOSEF TAMORY

Department of Biological Sciences
St. John’s University
Jamaica, NY 11439, U.S.A,

PATRICK LYNCH and BRIAN LOCKWOOD

Biscayne National Park
Homestead, FL 33090, U.S.A.

ABSTRACT

The vascular flora of Biscayne National Park, Florida consists of 372 species within 277
genera and 92 families. The a families in the flora are the Poaceae (47 species), Asteraceae
(34 species), and Fabaceae (28 species). The largest genera are Ti//andsia (7 species), Chamaesyce
(7 species), and Cyperus, Eugenia, Kalanchoe aot Pusiulum, each with 5 species. Each taxo-
nomic entry in the checklist is accompanied 2 an annotation, when appropriate, which

includes: C = cultivated species, not escaped; H = historical record, now extirpated; X =
naturalized exotic species; E = Florida eet T = Florida threatened; CE = Florida
commercially exploited.

Key worbs: flora, distribution, biodiversity, Biscayne National Park, Florida.

RESUMEN

La flora vascular del Biscayne National Park, Florida, esta formada por 372 especies
pertenecientes a 277 géneros y 92 familias. Las mayores familias de esta ae son las Poaceae
(47 especies), Asteraceae (34 especies), y Fabaceae (28 especies). Los ma s gén
Tillandsia (7 eae, ee a acne Cyperus, Eugenia, Klenow y aa con
5 especies cada uno. Cada entrada catalogo esta acompafiada de una anotacién,
cuando es plone que incluye: C = especie cultivada, no sstanpadias H = cita hist6rica,
rechazada ahora; X = naturalizada; E = en peligro en Florida; T = amenazada

en Florida; CE = eee loead comercialmente en Florida.
INTRODUCTION

Biscayne National Park, (BNP) comprising 181,000 acres /73,000 hect-
ares, is located between 25° 17' and 25° 40' N Latitude and 80° 05! and 80°
21' W Longitude (Fig. 1). The Park was designated in 1968 as a National
Monument, and, in 1980, the Monument was expanded and redesignated
as a National Park. The park is located approximately 20 miles (32 km)
south of Miami, Florida. The park’s northern boundary is near the southern
end of Key Biscayne, while the southern boundary is near the northern end
of Key Largo. Most of the land, (171,925 acres/69,577 ha) is submerged.

Stipa 18(4): 1207-1226. 1999

1208 Sipa 18(4)

|

Biscayne National Park

|| Mangroves & But tonwoods
WB Hardwoods
BB Grasses, Prairies, & Marshes

Fic. 1. Map of Biscayne National Park. Shown here are the plant communities referred to
in the flora.

The largest islands, listed in decreasing order of size are Elliott Key (1,651
acres/668 ha), Old Rhodes Key (641 acres/259 ha), Sands Key (416.7 acres/
169 ha), Totten Key (379.9 acres/154 ha), Little Totten Key (196.7 acres/
80 ha) and Long Arsenicker Key (124.9 acres/50.5 ha). There are 42 islands

—

in the park.

STALTER ET AL., Flora of Biscayne National Park 1209

The University of Georgia developed a map of major shoreline and up-
land vegetation units at the park in the summer of 1997 (Madden et al.
1999). These include mangrove (6905 acres/2794.4 hectares), hammock hard-
woods (126 acres/40.5 hectares), grasslands and marshes (197 acres/79.7
hectares). The total area of the shoreline and upland communities is 7416
acres/3001 hectares.

Climate

Climatological data presented is from the nearest weather station, Miami
Beach, 25 miles (40 km) north of the park headquarters. The climate of
Biscayne National Park is subtropical marine characterized by dry mild winters
and long warm summers (Jordan 1985; Winsberg 1990, 1992). Mean an-
nual temperature is 76.0° F (24.4 degrees C). January is the coolest month
with a mean temperature of 68.4° F (20.2° C) while August is the warmest
month averaging 83.1° F (28.4° C). Annual rainfall, generally a product of
convective sea breezes, is 46.02 inches (1169 mm). Occasional tropical storms
and hurricanes may produce heavy amounts of precipatation. Most of the
rain falls during the summer. August is the wettest month averaging 7.11
inches (180.6 mm) while March is the driest month averaging 1.56 inches
(39.6 mm). No frost was recorded at the Miami weather station between
1951 and 1974 (Anonymous 1974).

Detailed climatological data for Miami Beach and Tavernier, stations north
and south of Biscayne National Park, can also be found in the monthly publication
of the National Oceanographic and Atmospheric Administration. Temperature
data from 1980 to 1997 indicate that the temperature at Miami Beach reached
freezing on March 3, 1980 and on December 24, 1989. The lowest tem-
perature recorded at Tavenier was 35° F/1.1° C on December 25, 1989. A
freeze was recorded at the Turkey Point Weather Station, “for a few sec-
onds” on December 24, 1989 (personal communication with Ralph Heistand,
Florida Power and Light, Turkey Point Power Plant Meteorological Data
Manager) (see Fig. 2 for location of the Turkey Point weather Station).

Hurricanes

Hurricanes have ravaged Florida since time immemorial. Between 1900—
1989, Florida experienced 56 hurricanes. The strongest, a category 5 storm,
(see Fernald & Purdum (1992) for an explanation of Saffir/Simpson Hurri-
cane Categories) struck the Florida Keys September 3, 1935, killing 408
people. Other storms of note in the vicinity of BNP since 1960 were Donna
(1960), Betsy (1965), and most recently Andrew (1992). The major effects
of hurricanes on vegetation are wind damage and flooding associated with
the storm surge and/or heavy rainfall. Hurricane Andrew, a category 4 storm,
struck Elliott Key at 4:55 a.m. August 24, 1992. Hurricane Andrew's sus-
tained winds reached 230 kph (145 mph) with gusts in excess of 280 kph

1210 Sipa 18(4)

(175 mph) (Howard & Schokman 1995). Damage to vegetation on Elliott
Key was excessive. Twenty to thirty percent of the trees were downed and
large branches were sheared off almost all trees. Most small trees survived
with less structural damage, though many of the small trees were completely
defoliated. The trees at Adam’s Key, Biscayne National Park, were deunded
of leaves; most trees were broken off at 10—15 feet above the ground (Howard
& Schokman 1995).

Schinus terebinthifolius, a highly invasive exotic, suffered severe initial damage
from Hurricane Andrew. Many Schinus were blown down and nearly all were
defoliated. Individual §. terebinthifolius that survived the hurricane leafed
out more readily chan the native surviving mangroves. Howard and Schokman
(1995) observed that S. erebinthifolius appeared to have been spread by Hurricane
Andrew.

Native Rhizophora mangle fared poorly after Hurricane Andrew, especially
the dwarf forms, which were killed by the powerful storm. Avicennia germinans
and Laguncularia racemosa were also uprooted and killed by the hurricane,
although the surviving crees left standing sprouted soon after the hurricane
(Howard & Schokman 1995).

Water Salinit

Water quality nooo ey sites in the park are shown in Figure 2. Water
salinity ranged from 25.4 ppt at Turkey Point to 39.4 ppt at Caesar's Creek,
in 1997. There is little variation in water salinity at Alina’s Reef where salinity
ranged from 33.9 to 37.0. Salinity values are generally highest during June
when rainfall is low and evaporation high. Low salinity values have been
recorded near land during run-off immediately after heavy rains (Anony-
mows 199 >);

Historical Information
Prior to the Spanish cession of Florida to the United States in 1821, Native
Americans, Spaniards, Bahamians, and Americans established temporary
salvage camps in the Florida Keys (Leynes & Cullison 1998). The land presently
encompassing Biscayne National Park was sparsely settled in 1870; only
eighty five people were recorded in Dade County by the U.S. census at that
time (Niemec 1992). In 1877 Commodore Ralph Munroe reported that
there were “but a few dozen settlers,” in the Biscayne Bay area. The Florida
keys in the 1800s were valued for their mahogony. Because of the thin soil,
early settlers generally viewed the tropical key hammocks as “worthless”
for farming (Leynes & Cullison 1998). However settlers found that pine-
apple could be grown here, and by 1890 pineapple was well established as
a successful crop.

One of the earliest homesteaders on Elliott Key was Asa Sweeting, a Bahamian
immigrant, who moved to Key West in 1866. He emigrated to Elliott Key

STALTER ET AL., Flora of Biscayne National Park (21)

Biscayne National Park |

Turkey Point \
Turkey cout Channel \
sache Shoa if
riumph feel
aesars Creek
inas Reef

NOU RW

all Buoy

|
}

[> Long vid
Aw a: of”
p afl) . Key f ei]

% ni p : O Water Quality — — Mies.
| an ar Monitoring Sites Com?

if Kilometers

. Water quality monitoring stations and the Turkey Point meteorological station.
Sines Florida with the park boundary identifiec

from Key West in 1882, claiming 154.4 acres. The Sweeting holdings were
gradually enlarged to 238.8 acres. By 1887 he cleared and planted 30 acres,
gradually increasing his cultivated land to 100 acres. Pineapples and key
limes were raised as the main crops (Leynes & Cullison 1998), along with
bananas and tomatoes (Niemec 1992). A devastating hurricane in 1906

1212 Sipa 18(4)
signaled the end of profitable pineapple production. Key lime production
peaked in 1923 when more than 40,000 boxes were harvested. The destructive
hurricane of 1926 destroyed Sweeting’s lime groves, and the depression (1930s)
and competition with lime production in the West Indies and Mexico hin-
dered recovery. In 1935 lime production in the Florida Keys was a quarter
of the production of 1923. Moreover, the demand for other fruits and veg-
etables declined. By the late 1930s, sale of Elliott Key produce was nonex-
istent (Niemec 1992).

Development of land around Biscayne Bay and the Upper Florida Keys
grew rapidly as Miami Beach grew and prospered. The introduction of the
motorboat enabled wealthy individuals easy access to the nearby Keys, and
millionaires purchased the land on the Keys for weekend retreats (Leynes &
Cullison 1998). In 1904, Henry Flagler opened a fishing lodge on Soldier
Key. Carl Fisher established a vacation lodge on Adams Key in 1916, and
with several business associates, purchased Boca Chita Key as well. Dr. John
Gifford, subdivided and sold twenty-acre lots on Elliott Key stretching from
the bay to the ocean. Charles Brookfield operated a fishing camp, Ledbury
Lodge, on Elliore Key during the 1930s.

During the late 1950s, land “promoters” proposed to connect the Keys
to the mainland. After several years of wrangling over road construction,
Dade County officials decided against the construction of a causeway to the
islands, and stated, if a road were to be built, it would be financed by island
andowners. The island landowners incorporated, and created the City of
Islandia in 1960. Conservationists blocked Islandia’s efforts to build a causeway
to the islands, and in 1965 conservationists secured the support of the Hoover
Foundation to preserve the mea Keys. To foil the conservationists, the Islandia
governing officials approved the bulldozing of a highway, a 120'-wide strip
down the middle of Elliott Key, in the fall of 1967. The road, known as

“Spite Highway,” passed chrough a county park without authorization, and
destroyed 6.3 acres of vegetation. Remnants of this road persist today.

Congress approved the creation of Biscayne National Monument, and
President Johnson signed the bill into law on October 18, 1968. The bill
authorized the National Park Service to purchase and develop the park over
a five year period at the cost of twenty-five million dollars.

—

History of Floristic Work in the Park

Avery (1978) prepared an annotated list of 276 vascular plants on thirteen
keys of Biscayne National Park. Included on his list were frequency of oc-
currence (common, uncommon and rare, cultivated and non-native taxa)
and the distribution of the species on thirteen keys. The next study of im-
portance was that of Hammer and Bradley (1997) who reported 302 spe-
cies; no cultivated plants were included on their list. Four species, Spiranthes

STALTER ET AL., Flora of Biscayne National Park sali)

polyantha (Mesadenus polyanthus), Microg heterophylla, Neurodinm lanceolatum,
and Zamia pumila spp. pumila are known only from historical records. The
native status of six plants on the list prepared by Hammer and Bradley (1997)
is disputed: Acacia farnesiana, Carica papaya, Commelina diffusa, Cordia sebestena,
Occeoclades maculata and Yucca aloifolia. In addition, Lockwood et al. (1997)
prepared a list of the vascular plants at Biscayne National Park during the
summer of 1997.

Geology and Soils

Biscayne Bay is a shallow-water estuary along the southeast coast of Florida
ranging in depth from approximately one to three meters (Anonymous 1995).
An elongated ridge of Key Largo Limestone forms a border on the eastern
boundary of the park. Eleven bottom types are described from Biscayne Bay
including some spoil margins and mangrove soils (Anonymous 1995). The
soils on the Keys at Biscayne National Park are generally classified as Entisols
(Meyers & Ewell 1990). Soils are generally very shallow. The underlying
rock is porous Miami Oolite (Oolite limestone) of Pleistocene origin (Robertson
1955). Much of the rainfall is lost by runoff, or is leached through the po-
rous parent material (Stalter 1993).

METHODS

Collecting trips were made to the park in January, February, July and
December, 1997 and February, 1998. Objectives for each trip included the
collection of voucher specimens and accumulation of information on abun-
dance and apparent habitat preferences for each species.

More than 500 specimens collected above, herbarium specimens at Ev-
erglades National Park and Fairchild Tropical Garden, and plant lists gen-
erated by Avery (1978), Hammer and Bradley (1997) and Lockwood et al.
(1997), form the basis for this study. Taxonomically problematic specimens
were sent to various experts for identification; experts consultated include
Steve Clements (Amaranthaceae and Chenopodiaceae), Ihsan Al-Shehbaz
(Brassicaceae), Richard Mitchell (Polygonaceae), Robert Meyer (Poaceae),
James Montgomery (ferns), Charles Sheviak (Orchidaceae), Gordon Tucker
(Cyperaceae), Bruce Hansen, and Richard Wunderlin (various taxa). Voucher
specimens collected in the present study will be deposited in the herbarium
at Everglades National Park, Homestead, Florida, and partial duplicate sets
have been deposited in the herbaria of Brooklyn Botanic Gardens (BKL),
University of Michigan (MICH), Missouri Botanical Gardens (MO), New
York State Museum (NYS), University of South Florida (USF), James
Montgomery’s private herbarium (JM), and Fairchild Tropical Garden (FTG).
Accession numbers will be assigned by the staff at Everglades National Park
to the primary set of specimens at the herbarium at Homestead and will be

available upon request from the National Park Service.

1214 Stipa 18(4)

The annotated checklist contains an inventory of the vascular plants that
reproduce spontaneously and persist for more than one year without culti-
vation, and includes native taxa, naturalized and adventive weeds, escapes
from cultivation and historical records. In the checklist, plants are arranged
using the Cronquist (1988) system first by division, and then alphabeti-
cally by family and by species. Nomenclature follows Wunderlin (1998).
Synonyms are listed only for names used by Kartesz (1994).

RESULTS AND DISCUSSION

The vascular flora of the Biscayne National Park consists of 372 species
within 277 genera and 92 families. Ninety-three species, including nine
cultivated plants, or 25% of the flora, are not native to the region. A statis-
tical summary is given in Table 1, and an annotated list of species is in-
cluded at the end of the paper.

The major families include the Poaceae (47 species), Asteraceae (34 spe-
cies) and Fabaceae (28 species). Other large families are the Euphorbiaceae
(16 species) and Rubiaceae (13 species). Twenty-two percent of the species
comprising the total flora are contained in the Poaceae and Asteraceae. The
largest genera are Ti/landsia (7 species) and Chamaesyce (7 species).

Plant Communities

The flora of the Florida Keys has much in common with the West Indies
(Stalter 1993; Austin et al. 1987; Murphy & Lugo 1986; Correll & Correll
1982; Loope 1980; McGuire & Brown 1974; Thorhaug 1976; Robertson
1955). Davis (1942) recognizec

dune zone, scrub zone and hammock forest. Only the hardwood hammock

jon

four major communities: strand-beach zone,

community 1s an important community at BNP. Two additional commu-
nities, the mangrove forest and submerged aquatic communities exist at
Biscayne National Park.

The strand-beach community is dominated by halophytes (Stalter 1993).
Some species tolerate flooding, such as Cakile lanceolata and Sporobolus virginicus,
while others (e.g. Spartina patens) tolerate salt spray. Additional plants of
this community include: Chamaesyce spp., [pomoea pes-caprae, Panicum spp.,
Sesuvium maritimum, 8. portilacastrum and Aregusia gnaphalodes.

A poorly developed dune community exists in back of the strand-beach

—

zone. Common species here tolerate high concentrations of salt spray. Chamaesyce
mesembrianthemifolia, Spartina patens, Sporobolus sp., Suriana maritima, Argusia
gnaphalodes and Waltheria indica grow in this community. This community
is insignificant at BNP.

A xeric scrub ecotone, a transition zone of shrubs is poorly developed on
the Florida Keys (Davis 1942; Stalter 1993). Swriana maritima, Borrichia
arborescens, Lantana involucrata, Solanum bahamense and members of the Cactaceae

family are generally found here.

STALTER ET AL., Flora of Biscayne National Park

TABLE 1. Statistical summary of the vascular flora of Biscayne National Park, Florida.

Pteridophyta) Gymnospermae Magnoliopsida — Liliopsida Total

Families 5 l 70 16 92
Genera 8 | 195 56 277
Species 9 | 265 97 372
Native Species 8 | 206 64 288
Non-Native Species l 0 59 33 oe)

The hardwood hammock community contains a greater diversity of spe-
cies than the other plant communities. Many of the broad-leaved evergreen
trees that grow at BNP are typical of the Bahamas and Greater Antilles
(Davis 1942; Correll & Correll 1982; Fernald & Purdum 1992). Seven spe-
cies of andes are found in this community. Schinus terebinthifolius, a highly
invasive exotic is common here. S. terebinthifolius is a serious threat to na-
tive South Florida species, and is most abundant along the upland fringe of

the mangrove community. Some of the trees of this community are: Amyrzs

elemifera, Canella winterana, Coccoloba diversifolia, Coccoloba uvifera, Conocarpus
erectus, Cordia sebestena, Drypetes diversifolia, Eugenia foetida, Krugiodendyon ferrenm,
Lysiloma latisiliquum, Metopinm toxiferum, Panicum spp., Piscidia piscipula,
Pithecellobium spp., Schaefferia frutescens, Smilax auriculata, Suriana maritima,
Swietenia mahagoni, and Thrinax morrisii. For additional information on the
plant species comprising the hammock community, the reader is directed
to Harper (1927), Davis (1942), Forsgren and Kephant (1978), Robertson
(1955), Stalter (1993), and Myers and Ewell (1990).

Extensive mangrove forests are found in the intertidal zone. Mangroves
are sensitive to tidal flooding and are found in distinct zones. These spe-
cies, listed in order of their tolerance to tidal flooding are, RAbizophora mangle,
Avicennia germinans, and Laguncularia racemosa. Buttonwood, Conocarpus erecta,
is considered by Tomlinson (1986) as a mangrove associate, and is on the
upland fringe of the mangroves. Hurricane Andrew (1992) caused exten-
sive damage to this community, especially to RAzzophora mangle (Howard &
Schokman 1995). Extensive information on this community are in publi-
cations by Davis (1942), Tomlinson (1986) and Anonymous (1995).

The submerged “sea grass” community is the most extensive commu-
nity of the park comprising slightly more than 50% of the park. The sea
grass community is one of the most productive and important inshore marine
communities (Fernald & Purdum 1992; Myers & Ewell 1990). The “sea

rrasses” of importance are Turtle Grass, Thalassia testudinum, Manatee Grass,
Syringodium filiforme and Shoalweed Halodule wrightii. Sea grasses are dis-
tributed in accordance to their tolerance of salinity, and substrate prefer-

1216 Stipa 18(4)

ence (Fernald & Purdum 1992; Anonymous 1995). There is little salinity
variation in the bay near the islands where freshwater runoff during storms
is minimal (Fig. 2). Water salinity is more variable near the mainland when
runoff from torrential rains is substantial.

Several species in the flora are noteworthy because of their special status,
Psendophoenix sargentii, Sargent’s Palm, is believed to be planted in the Florida
Keys by early Bahamian settlers; it is not native to Florida according to
Avery (1978). Bruce Ledin in an unpublished manuscript disputed the na-
tive status of Sargent’s Palm in 1950. Avery (1978) reports that Aristolochia
pentandra, formerly known from Soldier Key is now restricted to Elliot Key.
Bucida buceras, Black Olive, and Spiranthus polyantha, Florida Keys Ladies
Tresses, were historically recorded on Elliot Key, though Avery (1978) be-
lieved that B. baceras was planted on Elliot Key. Kalanchoe-fedtschenko1, Lav-
ender Scallops, K. gastronis-bonniert, Palm Beachbells, and K. tabiflora, Chandelier
Plant were cultivated on Adams Key and persist after cultivation (Avery
1978). Clusia rosea was, “probably originally native to the Florida Keys,”
though at present it 1s only known from introductions (Wunderlin 1998).
Microgramma heterophylla was historically present on Adams Key, and is now
only extant at Key Largo (Avery 1978).

Three species, Schinus terebinthifolins, Casuarina equisetifolia, and C. glauca
are on the Florida prohibited plant Dist. 9 (DEP Chapter 62C-5 2). Coccoloba
uvifera is on Florida’s protected plant list. Plants, designated by the letter E
(Endangered) in the annotated checklist are on the State endangered list.

Species of foreign origin are only a minor component of the natural veg-
etation. Most non-native plants occur principally in ruderal sites, lawns,
and along the edges of trails and roads. Several exotics, Schinus terebinthifolius,
Casuarina equisetifolia and Colubrina asiatica pose a threat to the native spe-
cies in the park. The introduction of seventeen non-native species of grasses
(Poaceae), 18.3% of the non-native species, is especially noteworthy in the
flora. Other families containing a high number or high percent of non-na-
tive plants are the Agavaceae, Arecaceae, Crassulaceae, Fabaceae, and Sapotaceae.

ANNOTATED CHECKLIST OF SPECIES

The vascular plant taxa have been arranged according to the following
categories: ferns and fern allies, gymnosperms, dicots, and monocots. Within
each category, families and lower taxa are arranged alphabetically. Nomen-
clature primarily follows Wunderlin (1998) and Kartez (1994). Each entry
includes the following information sequence: native or non-native status
(C = cultivated species; not escaped, H = historical record, now extinct, X
= naturalized exotic species, E = Florida endangered, T = Florida threat-
ened, CE = Florida commercially exploited); scientific name; pertinent synonym,
enclosed in brackets.

on

STALTER ET AL.,

PTERIDOPHYTA

ADIANTACEAE/PTERIDACEAE—Maidenhair
a
ichum aureum L. Golden Leather Fern

Aoubann main Langsd.& Fisch.

siant
Preris vittata io Ge Ladder Brake; X
NEPHROLEPIDACEAE/DRYOPTERI-
DACEAE—Sword Fern Family
Nephrolepis biserrata (Sw.) Sehote uN. be ata
(Cav.) C. Christens.} Sword Fern; T

POLY PODIACEAE—Polypody Family
be heterophylla (L.) Wherry. Climb-
ing Vine Fern Historically present on
Key; now the only extant site
in Florida is on Key Largo. Hammer and

Nevrodium lanceolatum (L.) Fée, {Paltonium
lanceolatum (L.) K.Presl}. Ribbon Fern; E,H
Pleopeltis polypodioides var. michauxiana
Weath.) E.G. es & Windham,

P il) ipo. lium polyp fes (L.) Watt. var.

ee Weath. a Resurrection Fern

Ene ey

Whiskfern Family
L.) P. Beauv., Whisk-Fern
THELY PTERIDACEAE—Marsh Fern Family
Thelypteris Runthii li C.V. Morton,
Southern Shield Fer

GY MNOSPERMAE

ZAMIACEAE ily

Zamia pumila L. " (Zamia integr ee L.f.}
Florida eorioee Coontie;

Zamia F

MAGNOLIOPHYTA-
MAGNOLIOPSIDA

ANTHACEAE—Acanthus Family

Blechum pyramidatum (Lam.) Urb., Browne's
Blechum; X

ae cal (L.) Juss., Sixangle
Foldw

AIZOACEAE—
Sesuvinm maritimum (Walt.) Britton, Sterns,
Poggenb., Slender Seapurslane

Sesuvinm portulacastrum (L.) L., Shoreline

Seapurslane

AMARANTHACEAE—Amaranth IF

cee Family

Family

Flora of Biscayne National Park

1217

Alternanthera flavescens Kunth, Yellow
oywee

Amaranthus hybridis L., Slim Amaranth,
Pigwee

Blutaparon ernuculare (L.) Mears, Samphire;
Silverhead

Celosia nitida Vahl, West Indian Cock’s

Comb
Iresine diffusa Humb. & Bonpl. ex Willd.,
Juba’s Bush

NACARDIACEAE
eee 3 ‘a indica L.,
Metopium texiferum CL. ‘ Krug & Urb.,

oisonwoo

Schinus ee Raddi, Florida pro-

ibited plant; Brazilian Pepper; X

Cashew Family
70; X

ANNONACEAE—Custard- pone oy

Annona glabra L., Pond App

POCYNACEAE—Dogbane Famil

Catharanthus roseus (L.) G. Don, Madagascar

Periwinkle; X

Echites unbellata Jacq. Devils Potato;
Rubber

Nerium pee L., Oleander; C

Pentalinom ein (L.) B.F. Hansen &
Waunderlin, Wild Alamance Hammock

Viperstal
Rhabdadenia biflora Jacq.) Muell. Arg.,
Rubbervin angrovine

Thevetia pernviana (Pers.) K. Schum.,
Luckynut;
Vallesia antillana Woodson, Tearshrub; E

e

ARALIACE inseng Fam
anes sees (Endl ame Austra-
lian Umbrella Tree; Octopus Tree; X
ARISTOLOCHIACEAE—Birthworth Family
Aristolochia pentandra Jacq., Marsh’s Dutch-
man’s-Pipe; now restricted to Elliot Key;
formerly known from Soldier Key Tropi-
cal hammock’s; Rare; Dade County; E

ASCLEPIADACEAE—Milkweed Family
Cynanchum bahamense (Griseb.) Gillis,
Bahamium owwort; Bahama
Cynanchum
eres scoparium Nutt., Leafless
@ wor

c nif (Tac,

q.) Roem & Schult

White Twinevine; X

1218

ASTERACEAE—
Ambrosia avtenustifolia L.
, Perrenial Saltmarsh Aster

Aster Fami

I

, Common Ragweed
Aster tenuifolius L.
Baccharis angustifolia Michx., Saltwater

Falsewillow

Baccharis halimifolia L.., Groundsel Tree;
Sea Myrtle

Bidens alba (L.) DC. var. radiata (Sch. Bip.)
Ballard ex. T.E. Melchert, Beggarticks

Borrichia arborescens (L.) DC., Tree Seaside
Oxeye

Borrichia frutescens (L.) DC., Bushy Seaside
Oxeye

Chromolaena odorata (L.) R.M. King & H.
Rob., [Eupatorinm odoratum L.} Jack-in-
the-Bush

Complaya trilobata (1..) Strother, Creeping
Oxeye

Conyza canadensis (L.) Cronq. var. pusilla
(Nuce.) Cronquist, Dwarf Canada
Horseweed

Eclipta prostrata (L.) L.,

Emilia fosbergii Nicolson, Florida Tassel flower

Eupatorium eee (Lam.) Small,

False Daisy

ogfennel
Eupatorium leptophyllum DC., Falsefennel
Plaveria trinervia ee C. Mohr, Clus-
ered Yellow
fi pee linearis ae Neoware ‘llowtops

Penn-

rochaeriipy pensy vanicum WX/il Vai

avant Bverlaseing
Helianthus debilis Nutt.,
It Walter, Seacoast Marshelder

Duneflower
va imbricata
Launaea intybacea Jacq.) Beauv., Achicoria
zul;
Melanthera nived (L.) Small, [ML aspera Jacq.)
Small} Snow Squarestem
Mikania scandens (..) Willd.,

empvine

=f

Climbing

Partheninm hysterophorus L., Santa Maria
feverfew

Pectis glaucescens (Cass.) D.J. Keil, Sanddune
“inchweec

Pectis prostrata Cav., Spreading Cinchweed

Pluchea carolinensis Jacq.) D. aed
lobata (1..) Cass.} Cure-

Pluchea odorata (..) Cass., ee tscent

chea rosea R.K. Godfrey,

eed

Camphor
Solidago sempervirens L., Seaside Goldenrod

Rosey

“~
zy

Sipa 18(4)

Sonchus oleraceus L., Common Sowthistle; X
Tridax procumbens L., Coat Buttons; X
Vernonia cinerea (L.) Less., Little Ironweed; X
Wedelia trilobata (L.) A.S. Hitchc., creeping
oxeye
Youngia japonica (1...) DC., Oriental False
Hawksbeard; X
AVICENNIACEAE
Avicennia germinans (L.) L., Black Mangrove
BATACEAE—Saltwort Family
Batis maritima L., Saltwort; Turtleweed
BORAGINACEAE—Borage Family
Argusia gnaphalodes (L.) Heine, Sea Rose-
mary; Sea Lavender;
Bourreria succulenta Jacq., [B. ovata Miers |
ama Strongbark
Cor ‘lia sebestena L., Largeleaf Geigertree; C
He a ae angiospermum Murray,
orptonstail
ees CHVASSAVICHN 2 Seaside He-
liotrope; Salt Heliotrop
H Liotropin m polyphyllun ee m.
Ieliotrope
Tournefortia volubilts L., Twining So

, Pineland

dierbush
BRASSICACEAE—Mustard Family
Cakile lanc ae (Willd.) O. E

Coastal Searocket

Schulz,

Coronopus didymus (L..) Sm., Lesser Swinecress
Erucastrum eating )O.E. Schulz,
common Dogmustard
ee virginicum L., Virginia Pepperweed
BUDDLEJACEAE—Butterfly Bush Family
Polypremum procumbens L., Rustweed;
uniperleaf

family

BURSERACEAE
B ! eae: Gumbo-Limbo

CACTACEAE—Cactus Family
Acanthocereus tetragonus (L.) Hummelinck,
deena Ce actus; DUSOe Cactus; T
MVYISIGA Britton & Rose,
Prickly Arpplecacting R
Opuntia stricta Haw.) Haw., [O. stricta var.
dillenti (Xer-Gawl.) L.D.Benson] Erect
Pricklypear; T
CANELLACEAE—Canella Family
Canella winterana (L.) Gaertn.,

Pepper
Cinnamon; Cinnamon Bark; E

STALTER ET AL., Flora of Biscayne National

CAPPARACEAE—
eel ae L., Jamaican

Caper Family

o pee L., Bayleaf Capertree
CARICACEAE—Papaya Family
Carita papaya L., Papaya; X

CASUARINAC ice as ta Family
Cc ia L., Australian-Pine;

Hoievail Crustae Plofida prohibited
list;

Casuarina glauca Sieb. ex Spreng., Gray
Sheoak
Florida prohibited

CELASTRACEAE—Stafftree Family
Cros: Se rhacoma Crantz, Maidenbe rry;

; Suckering Australian Pine

—

ist; &

rae phyllanthoides Benth., Florida

yte

Ma
Schaefferia frutescens Jacq., Florida Boxwood, E
a arama CEAE
it lex hentandra
Saltbush
Salicornia bigelovii Torr., Annual G
DV Glasswort

Goosefoot Family

(Jacq.) Standl., Crested

—

asswort;

eige perennis Mills., {S. virgenica L.}
ial G wort

—

nia
Suaeda linearis (Elliott) Moq., Sea Blite;
Annual Seepweed
CLUSIACEAE—
Clusia rosea Jacq., Pitc
Bradley (1997) contend this taxon was,
“erroneously recorded as a native spe-
cies to the lower Florida Keys”, X

Mangosteen Fan
apple; ae and

COMBRETACEAE—Combretum Fami
Bucida buceras L., Black Olive. Records in-
dicate this was planted on Elliott Key;

that B. bucerus was
Hammer and

ee

“Tt is now believe
planted on the islands.”
Bradley (1997).

Conocarpus erectus L., Buttonwood
—. racemosa (L.) C.F. Gaertn.,
hite Mangrove
7 soa catappa L., West Indian Almond; X
CONVOLVULACE
Dichondra caroliniensis Michx.,

AE—Morningglory Family

Carolina
Ponysfoot

Ipomoea alba L., Moonflowers; Tropical
White enna lon)

Park 1219

Ipomoea indica (Burm.) Merr., Oceanblue
Morningglor
Ipomoea pes-caprae (L.) R. Br. spp. brasiliensis
(L.) Ooststr., Railroad Vine; Bayhops
Ipomoea violacea Heavenlyblue
Morningglory
CRASSULACEAE—Orpine family
Kalanchoe daigremontiana Raym.-Hamet &
H.Perrier, Devils Backbone; X
ease Hg Vis acs aym.-Hamet &
der Scallops; X
ne gastonis-bonniert Raym.-Hamet
alm Beachbells;
m.) Pers., Cathedral

er, Laven

& H.Perrier,
ahaneha

pinn sf we La

t; X
a . per ee ee ) Raym.-Hamet,
chandelier Plant; X
CUCURBITACEAE—Gourd Family
Melothria pendula L., Creeping Cucumber
Momordica charantia L., Balsampear, X
EUPHORBI EAE—Spurge Family
se a seein (Lam.) Muell. Arg.,
Bastard Cop
G humaese ie Engelm, ex, Hitchc.)
mestone Sandmat
Choma hirta (L.) Millsp., Pillpod

ra
Z

andm
Chama ‘icifolia (L.) Millsp., Graceful
Sandmat
Chama sopifolia (L.) Small, Hyssopleaf
{m

Choma mesembrianthemtfolia (Jacq.)
gand, Coastal Beach 8
C ee ophthalmica (Pers.)

Sandmat

D.G.Burch
Florida Hammock Sandmat

Chamaesyce porteriana Small Porter's Sandmat;

Drypetes diversifolia Krug & Urb., White-
wood; Milkbark; E

Drypetes lateriflora (Sw.) Krug & Urb., Guiana
Plum

Gymnanthes lucida Sw., Crabwood;
Oysterwooc

Hip tes fe tare L., Manchineel; E

Jatropha a ie Carils ush; X

Phy pe amarus Gakuen. & Thonn.,

jyale-of-Wind; Carry-Me Seed; X

Pointsettia cyathopbora (Murray) Bart

{Euphorbia cyathophora Murr.} Paintedleat

Fire-On-The-Mountain

Ricinus communis L., | ><

asain EAE— Pea Fa
ld farnesiana (L.) oa d.
er hs bonduc (L.) Roxb., Gray Nic
Caesalpinia major aes ) Dandy & Exe i
Hawait Pearls;
Canavalia rosea oa DC.,

, Sweet Acacia

Baybean; Sea-
side Jackbean
Crotalaria pumila Ortega Low Rattlebox
Dalbergia brownti Vacq. ) Schinz, Brown's
Indian Rosewood;
Dalbergia ecastophy len L.) Taub., Coinvine
Desmanthus virgatus (L.) Willd. » Wild Tancan
Desmodinm incannm DC., Vicker
Galactia striata (Jacq.) Ui. F lode a
mock Milkpea
Gliricidia sepium (Jacq.) Kunth ex Walp.; X
Indigofera spicata Forssk., Trailing Indigo;

Leucaena lencocephala (Lam.) de Wit, White
eadtree; X

Lysiloma latisiliquum Benth., False Tama-
rind

Medicago lupulina ee Black Medick; X

Melrotis albus Medick., White Sweetclover

Melilotis officinalis a am., Yellow
Sweetclover;

Parkinsonia acyleat.

rusalem Tho
Pix i i aie Seep. I
e; Jamaican Dogwe
ee in dulce ene ‘ un

., Mexican Palo Verde;

a Fishpoison

.. Mon-
od;

Photon as Britton ex Britton &

e, Florida Keys Blackbead; T
, Catclaw

Lose,
ae isi -cati (L.) Benth.
Blackbea

Rhynchosta minima (1..) DC., Least Snoutbean

Sophora tomentosa 1.., Yellow Necklace

Stylosanthes hamata (L.) Taub., C ee

Tamarindus indica L., Tamarind; X
Trifolium repens L., White Clover; Dutch

ver;
Viena luteola Jacq.) Benth.,
Lowpea
GENTIANACEAE— Gentian Family
Enstoma exaltatum (L.) Salisb. ex G. Don,
Marshgentian; Catchfly; Prairie gentian

Hairypod

Stipa 18(4)

Leiphaimos parasitica Schltdl. & Cham.,

Voyria parasitica (Schltdl. & Cham.)

Ruyters. & Maas.]} Parasitic Ghostplant; E
HALORAGACEAE— —Watermilf oil family

rsh Mermaidweed

HIPPOCRATEACEAE—Hippocratea Family
Hippocratea volubilis L., Medicine Vine
LAMIACEAE—Mint Family
Salvia serotina L., Little Woman
LAURACEAE—Laurel Family
Cassytha fliformis L., Love Vine; Devil's Gut
Ocotea coriacea (Sw.) Britton, Lancewood
LENTIBULARIACEAE—Bladderwort Family
Utricularia foliosa L., Leafy Bladderwort

LOGANIACEAE—1L ogania Family
Mitreola petiolata (J.F. Gmel.) Torr. & A.Gray,
[Cynoctonum mitreola (L.) Britton} Lax
Hornpod
LYTHRACEAE—Loosestrife Famil
Ammannia latifolia L., Pink aes
Toot theup
MALVACEAE—Mallow Family
Abutilon permolle (W/illd.) Sweet, Coasta!
Indian Mallow
(L.) Brizicky, B

var. rosd-Sinensts,

laddermallow

Hibiscus rosa-sinensis L. v
semallow; Shoe-Back-Plant, X
Hibiscus tiltaceus L.., Sea Hibiscus; Mahoe; X
Matvastrum corchorifolium (Desr.) Binion
ex Small, False M
Sida abutifolia P. Mill, Spreading Fanpetals
Common Wireweed;

Mallow

Sida acuta Burm. f.,
Common Fanpeta

Sida ciliaris L. “Bracted Fanpetals; Fringed
Fanpetals

Sida rhombifolia .., Caban Jute; Indian
Hemp

Php pp .) Sol. ex Correa, Portia

MELIACEAE—Mahogany Family
Swietenia mabagoni (L.) Jacq., We st Indian
Mahaogany; E

MORAC

Mulberry Family
Ficus altissima Blume, Counsil Tree; X
Ficus aurea Nutt., Strangler Fig; Golden Fig
Ficus citrifolia Mill., Wild Banyan Tree
MYRSINACEAE—Myrsine Family

STALTER ET AL.,
Ardisia escallonioides Schiede & Deppe ex
Schldl. & Cham., Marlberry

MYRTACEA
Cal che ae Grisel
icew

E—Mpyrtle Family
4 Pale I idflower;

ae pe (Sw.) Bes , White stopper
Eugenia confusa DC., erry Stopper
Eugenia foetida ae
Boxleaf Stopper
Eugenia rhombea Krug & Urb. ex Urb., Red

ane Stopper;

Stopper
Eugenia uniflora L., Surinam Cherry; X
NYCTAGINACEAE—Four O'Clock Family
Boerhavia diffusa L., Red Spiderling;

)

Boerhavia erecta L., Erect Spiderling

Bougainvillea ebes cme Paper Flower;
».4

Guapira discolor (Spreng.) Little, Beeftree
Pisonta aculeata V.., Devilsclaws; Pullback
OLACACEAE— Ximenia Family
Ximenia americana L., Tallow Wood; Hog Plum
—Olive Family
; san sambac(L.) Aiton, Arabian Jasmin;

ONAGRACEAE—Evening Primrose Family
Gaura angustifolia Michx., Southern
Beeblossom
Ludwigia b {. Micheli) H. Hara,
Crelns Pianos Willow
Ludwi — actovalvis Jacq.) Raven, Mexi-
an Primrose-Willow
OX ALIDACEAE—Woodsorrel Family
Oxalis corniculata mon Yellow

*?
Woodsorrel; Creeping Woodsorrel

PAPAVERACEAE—Poppy Fam
— mexicana L., Mexican au Ae Poppy
PASSIFLORACEAE- Passion-Flower Family

Pasifloa multiflora L., Whitetlower Pas-
sion-Flower; E
Passiflora suberosa L., Corkystem Passion-

PHYTOLACCACEAE—Pokeweed Family
Rivina humilis L., Rougeplant
PLUMBAGINACEAE— y
Limonium carolinianum (Walter) Britton,
Carolina Sealavender

Leadwort Famil

Flora of Biscayne National Park 122),

POLYGON ACEAE—Buckwheat F
Caccoloba diversifolia Jacq. , Tee es

ere uvifera (L.) L., Seagrape. Protected
by Florida Law 370.041
PORTULACACEAE—Purslane Family
Portulaca oleracea L.., Little Hogweed; X
Portulaca pilosa L., Pink Purselane; Kiss-
Me-Not
PRIMULACEAE—Primrose Family
Samolus ebracteatus Kanth, Water Pimpernal
nolus valerandi L. subsp. parviflorus (Raf.)

>=
Ss
aa

Hultén, Pineland Pimpernel; Seaside

Brookweed

RHAMNACEAE—
Colubrina arborescens (Mill.) Sarg., Greenhart;

Buckthorn Family

E
a astatica (L.) a. Leather-
lea n Nakedwood; X
a. se (Sw) Briziky. & W.L. Stern,
Soldierwood;
Gouania lupuloides (L.) Urb., Chewstick;
Whiteroot
Krugtodendron ferreum, (Vahl) Urb., Black
Ironwood; Leadwood
Reynosia septentrionalis Urb.,
RHIZOPHORAC
Rhizophora mangle L., Red ues ale

Darlingplum;’'T

Mangrove Family

RUBIACEAE—Madder Fan

Chiococta alba (L.) ie {C. parvifolia
Wullschl. ex Griseb.} Snowberry;
Millebes

Exostema cart 2 Jacq.) Schult., Carribean
Peace &

Gardenia angusta (L.) Merr., Gardenia; C

Genipa clusitfolia Jacq.) Griseb., [Casasia
clusiifolia Jacq.) Urb.] Sevenyear Apple

Guettarda elliptica Sw., Hammock Velvetseed

Guettarda scabra (L.) Vent., Rough

elvetseec
Hedyotis corymbosa (L.) Lam., {[Oldenlandia
corymbosa L.} Flattop Mille Graines
Morinda royoc L., Redgal
Psychotria nervosa ape Wild Coffee
Randia aculeata L., White Indigoberry
he i (Cham.& Schltdl.)
& Schult. f., Largeflower Mexican
ae

Spermacoce floridana Urb., Florida False

Buttonweec
Spermacoce verticillata L.,

Shrubby False

Buttonweed;
RUTACEAE—Cicrus Family
cad as L., Sea Torchwood

tifolia (Christm )Swingle

, Key

Lime
Citrus aurantium L., Sour Orange; C
Citrus limon (L.) Burm. f., Lemon; C
Citrus reticulata Blanco, Tangerine; C
Zanthoxylum fagara (L.) Sarg., Wild Lime;
Lime Pricklyash

SAPINDACEAE—Soapberry Fam
Cardiospermum haticacabum 1. te in-a-
Puff;

Exothea paniculata Juss.) Radlk. ex Durand,
I Beescouet 1
[7 bij acq., Spanish Lime; X

y Jac
ore ere L., Soapberry
SAPOTACEAE—Sapodilla Family
ChrysophyHum olivifor meL.,S Satinleaf: T
Manitlkara jaimiqui (C.Wright ex Griseb.)
Dubard ssp. ewarginata (L.) Cronquist,
Wild Dilly
Manilkara zapota (L.) P. Royen, Sapodilla; X
Pouteria ssl (Kunth.) Baehni, Egg
Fruic, Canistel;
anes omingensts — f.) Baehni; X
a i T.D. Penn.,
Saffron
Sideroxylon foetidissimun Jacq., False Mastic
Sideroxy ie salicifolinm (L.) Lam, , Willow
Bustic; White Bully

spr

SCROPHULARIACEAE—Figwort Family
/ esiuaaain - tb-of-Grace

ene ld oie L., Goatw
Linarta canadensis L. Chaz., . uttallanthus
canadensis (L.) D.A. Sutton} Canada
Toadflax
SIMAROUBACEAE—Quassia Family
Simarouba glauca DC., Paradisetree
oe ne EAE oe rade Family
var. glaby oe — )
rd Pepp

ADT,

He ser & saree
oo Pey eae
| m Walter, Chriscmasberry;

Carolina Desertthorn

Stipa 18(4)

—

rnericaniumn P. Mill.. American Black
ightshade

Nightsl
Solanum bahamense L.,

Syl ay

1 }))

Bahama Nightshade
Solanum chenopodioides Lam., {S. nigrescens
Martens & Ga/eotti misapplied}, Black
Nightshade
Solanum evianthum D. Don, Potatotree
STERCUL i Cacao Family (Includes
Byttne
W ae Ld ve L.
URIANACEAE— Bay-Cedar Family
Suriana maritima \., Bay

, Sleepy Morning

THEOPHRASTACEAE—Theophrasta Family
cue keyensis Mez, Joewood; T
ULMACEAE—E]Im Family
Trema jae ies (Schult.) Blume, Pain-
in-che- » Wese Indian Trem
U pan AC e oe Siare Family
Nutc., F fer a Pellitory
Hanna ais )Liebm.., Artille Ty Plant;

Rockweed

VE a NAC et —Vervain ae
itha pi tL. {C. fruticosum L.}

EF lorida Piddlewood
Lantana camara \.., Lantana; Shrub Ver-
bena; X
a involucrata L., Buttonsage
Phyla nodiflora (1..) Greene, Capeweed;
ac Tangle Eo rfruit
Stachytarpheta jamarcensis (L.) Vahl, Blue
Porterweed; Joce
VITACEAE—Grape Family

Vitis rotundifolia Michx., Muscadine

=

ZYGOPHYLLACEAE—Caltrop Fami
Guayacum sanctum L.., Holywood Lignumvyitae;

EF
y

MAGNOLIOPHYTA-LILIOPSIDA
AGAVACEAE—Agave Family
Agave decipiens Baker, False Sisal
Agave sisalana Perrine, Sisal Hemp; X
Alpes vera L., Aloe; X
Sansevieria hyacinthoides (L.) Druce, Bow-
string Hemp; Mocther-in-Law-Tongue,;
African Bowstring;
Sansevieria trifasciata Prain cv. ‘Laurentit’,

Bowstring Hemp;
Yucca aloifolia L., Spanish Bayonet; Aloe

STALTER ET AL., Flora of Biscayne National Park

Yucca; X
ALISMATACEAE
Sagittaria Acai Michx. var. weatherbiana
(Fe

—Water-plantain family

rnald) Bogin, Weatherby’s Arrowhead
Sagittaria pen L., Bull Tongue
Arrowheac
AMARYLLIDACEAE—Amaryllis eee
Hymenocallis latifolia (Mill.) M. Roem.,
Mangrove Spiderlily; een iSpidedily
ARACEAE—Arum Family
Epipremnium pinnatum (L.) Engl., Golden
een eae yee SS
ta (Jacq.) L.H. Bailey,
_ Florida a Palm; T
acos nucifera L., Coconut Palm
ae datty, a. Date Palm; a Fruit;

X

Phoenix reclinata Jacq., Senegal Date Palm; X

Psauaophoenin aes Z Wendl. ex Sarg.,

argent’s Cher

Sabal cae pale ‘Lod. ex Schule.
& Schult. f., Cabbage palm

Thrinax morrisii H. Wendl., ag Th

m; Key Thatch Palm

Pipe radiata Lodd. ex os & Schule.
fs, ee ba sali auct. non Sw.} Florida
Tl Palm;

Veitchia seers a H.E. Moore, Ma-
nila Palm; X

atch

lool

B3BROMELIAC —Pineapple Family
Vilas A Schule. & Schult. f.,
Nor n Needleleaf, T
Z ints, eek wlata Sw., Cardinal Airplanc;

L i flexuosa Sw., Twisted Airplant; E
i ilandsia pauifolia Baker, aya Airplant
Tillandsia recurvata Ballmoss
Tillandsia nsneoides (L. ) L. , Spanien Moss
#ta L., Giant Airplant; E

wndsia utvicit:

COMMELINACEAE—

Spiderwort Family

Commelina diffusa Burm. f., Dayflower

aa ie ae (Rose) D.R. Hunt,
rplequeen;

Tada ee Sw., [Rhoeo ae
oses-in-the-Cra
ee ee x

CYMODOCEACEAE—Manateegrass Family

by

1223

Hatlodule wrightii Ashe, (H. beaudette (den
) den Hartog} Shoalweed
Syringodium filiforme Kutz., [Cymodocea
filiformis Kutz.) Correll] Manateegrass
CY PERACEAE—Sedge Family
Cladinm aces se Crantz, [C. mariscus (L.)
ae ia aaa Kuk.]}
Jamaica oo weras
a esculentus L., cates —— Chufa
Flatsedge
Cyperus involucratus Rottb., Umbrella Plant; X
Cyperus ligularis L., Swamp Flatsedge
sie iantolies L.C. Rich., Flatleaf
edg re
a ea Rottb., Manyspike

Flatsegde
Eleocharis equisetoides (Elliott

) Torr., Jointed
Spikerush
Rhynchospora colorata (L.) H. Pfeiff., Starrush
Whitetop

HY DROCHARITACEAE—Frog’s Bit Family
aeicaue pee Ostenf., Carribean

era
Thalassia ae Banks & Sol. ex. J.
Koenig, Turt
LILIACEAE—Lily
Amaryllidaceae
MUSACEAE—Banana Family
Musa acuminata Colla, ‘Dwarf Cavendish’
Dwarf Banana;

ae

egrass

Family, See Agavaceae and

ORCHIDACEAE—Orchid Family
Bletia oe (Lam.) DC. [O/etis ee

am.) A. de Candolle} Pine P
Encyclia oe (Lindl.) eee r, Dols
lar Orchid; E
Eng ee (Lind] .) Small, Florida
y Orchid;
rete mat (Lind1.) Lindl., Monk

Se a antha Rchb. f., Florida Keys
Ladies Tresses; Gray Pesce. His-
torically reported from Elliott Key;

POACEAE—Grass Family

Andropogon oF ee Britton,
Ster ggenb., Bushy Blueste

ig oniven pee Trin. ex rl

Beardgrass

~~
~~)

Andropogon virginicus L., Brownsedge
Bluestem

Bothriochloa barbinodis (Lag.) Herter,
Beardgrass; X

Bothriochloa pertu samus,
{Andropogon se 7 ) aie Pikied
Beardgrass;

Roem. & Schult
Slimbristle Sandbur

Cenchrus browntt

Cenchrus echinatus L., Southern Sandbur

tis, [(C. carolinianus
Walt. } Coastal Sa:

Cynodon dactylon (L.) Pers.,
X

Cenchrus incertus M.A. Cu
Bermudag rass;

TD. tylo pyr tspy)

yptium (L.) Willd. ex Asch.
‘Dorban Crowfootgrass; X

& Schweinf.

Digitaria bicornis (Lam.) Roem. & Schult.
ex. Loudon, Asia Crabgrass; X

Digitaria ciliaris (Retz.) Koeler, Southern
Crabgrass

Digitaria filiformis (L.) Koeler var.
dolichophyHa (Aenrard) Wiptt, Carribean
Crabgrass

Distichlis spicata (L.) Greene, Sa

Elensine indica (L.) Gaertn.,
Goosegrass;

Eragrostis amabilis (L.) Wight & Arn., LE.
tenella (L.) P. Beauv. ex Roem. & Schult. ]
Feather Lovegrass,; X

Eragrostis ciliaris (L.) R. Br.,
Lovegrass;

Eragrostis elliottii S. Watson, LE. campestris
Trin.} Ellioct’s Lovegrass

Eusta by § petraca (Sw.) Desv., [Chloris petraca

uneepees Fingergrass

T acpzarcye Ar4 ta (L.)

Florida vi

terass
Indian

Gophertail

Hitchc., Smallcane:

Monanthochloe littoralis Engelm., Shoregrass;
Keygrass

Neyraudia i eaaiee (Sunth) Keng ex
it

(a)

ic., Burmareed; Silkreed; X
Panicum amarum Elliott, Bitter Panicgrass
Panicum repens L. Pop ass; X
ae HM VIN: Lata L., Swite

lt, ettit Chapm., Coral Paspalum;

Sp
Laden S C rowngrass

” L., Knotg

Paspalum notatum Flue gee, Bahiagrass; X

Paspalum distichun

SMILACACEAE—

Sipa 18(4)

Paspalum setaceum Michx., Thin Paspalum

Paspalum peat Sw., Sandlot Paspalum

Pennisetum polystachion (L.) Schult.,

sain setosum (Sw.) Rich] West
dian Pennisetum; Missiongrass

Rhynchelytrum om Wi Id.) C.E. Hubb.,

Natalgrass; X

_

Schizachyrinm sanguineum (Retz.) Alston,
{S. semiberbe Nees] Crimson Bluestem

Schizachyrium scoparinm (Michx.) Nash, Little
Blustem

Setaria herlyrttol rea ( fon Stapf. P: ASC
taria palma (J. Konig) Stapf, Palmgrass,

Setavia parviflora (Poir.) Kerguelen, [S.
geniculata (Poir.) P. Beauv.] Yellow
Bristlegrass; Knotroot Foxtail

oe patens (Aiton) Muhl.,

rdg rass; S : Saltmeadow Cordgrass

Spor robolus dom is (Trin.) Kunth, Coral
Dropseed

Sporobolus indicus (L.) R. Br., [S. posrettsz
Roem.& Schult.) Hitche.] Smutgrass

Sporobolus pyramidatis (Lam.) Hitchce.,
Whorled Dropseed

Sporobolus virginicus (L.) Kunth, Seashore

Marshhay

jos

45]

Dropseed

Stenotaphrum secundatum (Walter) Kuntze,
St. Augustinegrass

Urochloa adspersa (Trin.) R.D. Webster,

achiavia adspersa (Trin.) Parodi]

Dominican Signalgrass

Urochloa mutica (Forssk.) Nguyen,
{Brachiaria mutica (Forssk.) Stapf.},

Paragrass; X
Urochloa subquadripara (Trin.) R.D. Webster,
{Brac taria subquadripara (Yrin.) Hitche.}
Tropical Signalgrass; X
Loystat ee Steud., Korean Templegrass;
Japanese
Zoysia
emplegrass; X

Lawngrass;

matrella (L.) Merr., Manila

Smilax Fam

TY PHACEAE—

Smilax auviculata Walter, at af Greenbrier
Smilax havanensis Jacq., Everglades Greenbrier

Cattail Fami

<

Typha domingensis Pers., Southern Cattail

STALTER ET AL., Flora of Biscayne National Park l.

NO
No
“A

ACKNOWLEDGMENTS

Appreciation is expressed to Superintendent Richard Frost for unrestricted
access to Biscayne National Park. We thank Everglades National Park and
Lynka Woodbury, Fairchild Tropical Garden, for access to their herbaria.
There were many others with the National Park Service, too numerous to
mention here, who helped us in so many ways; we thank them all. For as-
sistance in the identification of several taxa we thank Ihsan Al-Shehbaz, Steven
Clemants, K.N. Gandhi, Bruce Hansen, Robert Meyer, Richard Mitchell,
Jim Montgomery, Richard Rabeler, Charles Sheviak, Gordon Tucker and
Richard Wunderlin. For GIS maps we thank Barry Wood. Finally, we ac-
knowledge the assistance of Ali Pootrakul, and Andrea Arsene research students
at St. John’s University, and the financial support of St. John’s University.

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Austin, D.FE, A.F CHoLewa, R. B. Lasrrer, and B.F. Hansen. wet The Florida of John
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Avery, G.N. 1978. Biscayne National Monument plant distribution and frequency list.
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CORRELL, B.C. and H.B. Correit. 1982. Flora of the Bahama Archipelago. Lubrect and

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Gut ist, A. 1988. The evolution and ¢

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University Press of Florida,

A CHECKLIST FOR THE VASCULAR PLANTS
OF PADRE ISLAND NATIONAL SEASHORE

[Ge ee A.D. NELSON’, J.R. oo
MACKF?, T. WILBURN’, and A. DA

'Department of Biology, Box 158
Texas AGM University-Kingsville
Kingsville, TX 78303, U.S.A.
°Department of Biological Sciences, Box T-O100
Tarleton State University
Stephenville, TX 76402, U.S.A.
Science Department
Laredo Community College
Laredo, TX 78040, U.S.A.

ABSTRACT

The flowering plants of Padre Island National Seashore (PINS) were sampled over a two
year period in three different regions of the island. Specimens from our investigation and
plants reported for PINS from the literature were compiled into a checklist. The flora con-
tains both Flaveria brownii and Sporobolus tharpii which are endemics of the western Gulf of
Mexico barrier islands. Lyciwm carolinianum var. quadrifidum, reported only along bayshores
in Texas, was found in the tidal flat community. Two hundred and we nine genera and
456 species of flowering plants were identified as part of the PINS flora. These occur in

families with the Poaceae (95 spp.), Asteraceae (57 spp.), Fabaceae (44 and Cee
(29 spp.) most common.

RESUMEN

Se han tomado muestras de las angiospermas del parque nacional “Padre Island National
Seashore” (PINS) durante un perfodo de dos afios, en tres dreas diferentes de la isla. Se hizo
un catdlogo de los “especfmenes” de nuestra een y de las plantas citadas a PINS
procedentes de una serie de articulos. La flora contiene tanto Flaveria or owni? COMO os obolus
tharp1i que son endémicas de las islas de os del Oeste en el Golfo de Méxic ha
encontrado en la comunidad de la zona mareal plana Sai carolinianum vat. sna ea
que antes solamente se habfa citado a lo largo < las bahfas de 259 gér 456
especies de anglosper mas fueron ecco en la flora de PINS. Las aie mencionadas

pertenecen a 77 familias siendo las familias Poaceae (95 especies), Asteraceae (57 esp.), Fabaceae
(44 esp.), y Cyperaceae (29 esp.) las mas ec,
INTRODUCTION

The Texas Coast is framed by sandy barrier islands which are floristically
similar to other barrier islands of the Gulf of Mexico and Atlantic Coast.
However, upland sites on Texas barrier islands often grade into extensive

Sipa 18(4): 1227-1245. 1999

1228 Stpa 18(4)

grassland areas (Duncan & Duncan 1987). Padre Island National Seashore
(PINS) encompasses most of the barrier island often referred to as northern
Padre Island (Figure 1) and contains extensive grassland areas between its
foredune ridges on the eastern gulf side and tidal flats bordering the west-
ern Laguna Madre. Southern Padre Island has been considered separately
from northern Padre Island in terms of floristic analysis (Judd et al. 1977;
Lonard et al. 1978; Lonard & Judd 1980) and is physically separated by the
Mansfield Ship Channel. In contrast, Corpus Christi Pass once separated
Padre Island from Mustang Island but this pass is now filled and the two
islands are joined (Weise & White 1980). There have been a number of
investigations involving the vegetation of saa and wetland habitats on
northern Padre Island. Most recently, Nelson et al. (1997, 1999) examined
four communities in regard to vegetational ecology and floristics near Big
Ball Hill on PINS (Fig. 1). Drawe and Ortega (1996) examined effects of
seismic survey vehicles on vegetation from five different communities on
the northern one-third of the island during October 1988 as well as April
and September, 1989. Also, Brown et al. (1989) compiled a vegetative his-
tory of the island summarizing early historical accounts and scientific works.
Carls et al. (1991) assessed the impact of oil industry activities on three
plant communities on the northern one-third of the island, and McAtee
and Drawe (1975, 1980, 1981) examined the impacts of vehicular and pe-
destrian traffic patterns on the northern one-third of the island. However,
the most comprehensive ecological treatment of PINS vegetation was done
by Baccus and Horton (1979) and Baccus et al. (1977). They examined vegetation
along the entire length of PINS and focused on areas with differing amounts
of vehicular and pedestrian traffic. Drawe et al. (1981) and Kattner (1973)
conducted comprehensive vegetation studies of five habitats about 13 km
south of the PINS Ranger Station during each of six seasons from winter
1972 tospring 1973. Higginbotham (1972) analyzed vegetation data from
four habitats in che northern one-third of the National Seashore, and Orteni
etal. (1972) investigated dune vegetation near the PINS Ranger Headquarters.

PINS was established in 1963 and since that time a considerable num-
ber of ecological investigations have been conducted, whereas only four floristic
studies have been completed. Rabalais (1975) produced a list of the flora of
PINS using work by Jones et al. (1961), Jones (1966), and Rechenthin and
Passey (1967). A checklist incorporating data from the aforementioned ecological
and floristic studies has never been published. The purpose of this investi-
gation was to produce a checklist for PINS using data from the literature as
well as our own field studies of the island. A checklist can aid researchers in

identifying plants, provide data for the development of a flora or manual,
and allow for biogeographic comparisons with other islands (Nelson Goetze
& Lucksinger, unpublished data).

NeGreteE ET AL., Vascular plants of Padre Island National Seashore 1229
METHODS AND MATERIALS

Areas known as Big Ball Hill, Ranger Road, and Bird Island Basin were
sampled during the investigation (Fig. 1). In the Big Ball Hill region, floristic
surveys were conducted in May and August through November of 1996, as
well as January through March and May of 1997. The Ranger Road and
Bird Island Basin regions were sampled from January through April of 1998.
Sampling was done by establishing approximately 500m belt transects in
each community type with the aid of a Magellan GIS navigator. Commu-
nities identified for sampling included coppice dunes, foredunes, barrier
flats and secondary dunes, tidal flats (Nelson et al. 1998), and disturbed
areas such as roadside ditches. Transects in each community were walked
and plants in flower within sight of the transect were sampled. In a few
instances, woody vegetation was sampled that was not in flower. Plants collected
along the transects were identified by using Jones (1982), Correll and Johnston
(1970), Gould (1975), and Gould and Box (1965). Species nomenclature
was updated to Jones et al. (1997). Common names were taken from Correll
and Johnston (1970) taking into account nomenclatural changes based on
Jones et al. (1997). Vouchers were deposited in the herbarium at Texas A&M
University-Kingsville (TAIC) and Tarleton State University (TAC) in
Stephenville, Texas. In addition to our field data, information from the in-
vestigations summarized in the introduction were used in compiling the
checklist. Rechenthin and Passey (1967) reported several specimens only
as genera; these records were not included in our checklist. These include
Allium, Abutilon, Callirhoe, Desmodium, Penstemon, Polygonum, Rumex, and Tragia.
The checklist is arranged alphabetically by and within family and includes
the scientific name, authority, common names (if available), herbarium, collector
(if available), and collection number. Acronyms used for Texas herbaria in-
clude: C.C. Museum (Corpus Christi Museum), PINS (Padre Island Na-
tional Seashore, Corpus Christi), TAC, and TAIC. When collection data are
not available, the abbreviation NSA (No specimen available) is used and
investigations where this species was reported are listed. Following the collection
data are annotations on habit, endemism, and habitat. Plants that are woody
are listed as such; plants without this designation are herbaceous in habit.
When known, habitat designations of coppice dunes (CD), foredunes (FD),
barrier flat (BF), and tidal flat (TF) (Nelson et al. 1997, 1999) are used.

—

RESULTS AND DISCUSSION
Seventy-seven families, 259 genera, and 456 species are repoted from PINS.
The five most common families are Poaceae (20.8%), Asteraceae (12.5%),
Fabaceae (9.6%), Cyperaceae (6.4%), and the Euphorbiaceae (4.2%). The
flora of PINS primarily represents a subset of the flora of the South Texas

)

~

Sipa 18(4)

Gregory
Portland fai} ‘Acanaos Pass
HA
a ‘Aronses Pass
EER os Pret Aronsos

Pw
4
fs
oS
yy
S

4M,

LY, Water Exchange Poss

x
’ iN
<4 ‘up,
~ o
=
Ss Re

NUECES CO.

AEEDERG co. § o Park Rd. 22
OLY
‘ay

2

2
LIX

agers

Bird Island Basin Study Site
Ranger Road Study Site

Sragste,
eet

04

q
qn f] N
, SS: ; ; .
AY oCPKLEBERG RN Big Ball Hill Study Site
O° | KENE DCO” NSP :
N $
0 sayy
mRSs a
1 ey s
& ;
RSS =~
1 kN 8
a | oN re
nl BN
INNS %
( iP RN * Bolivar Penintulo
18 [EX a Golveston isiond
Pig SN = Foltats Istond
I RS Matagorda Peninsulg co
18 AS 2 Matagorda Intand et !
13 Nh 1. Joseph Island w
s/s NY 7 juslang Island
1s \
od 18 NN v
Ios NN 3
io
3 NY
oe
Lay — Oo 25 50m
SS ame ane)
NAS > 40 60k
\ RS 2
RSNEOY GO: Ra
WILLACY CO Sr SN
N
South Padre Islond
Gores Sontic
Poss mv 9 5 10 mi
Brozos Islond }/+—_ +
io) 8 16 km

Fic. |. Map (Weise & White 1980) indicating the location of the study sites. Hatched
lines indicate the boundaries of Padre Island National Seashore. Inset shows location of

Padre Island National Seashore in Texas.

NEGRETE ET AL., Vascular plants of Padre Island National Seashore hood

mainland with the exception of Flaveria brownii and Sporobolus tharpit, which
are reported as possible endemics on Padre Island and the Tamaulipan bar-
riet islands of northern Mexico (Britton & Morton 1989). Lycium carolinianum
var. guadrifidum, reported only along bayshores in Texas (Britton & Morton
1989), was found in the tidal flat community of PIN

The lack of woody vegetation is a striking feature in regard to the PINS
flora. Less than 10% of the species from PINS are considered subshrubs,
shrubs, or trees. When one considers the populations of these species in comparison
to the total vegetative community of the island, it is estimated that less
than 1.0% of the vegetation is woody. Based on historical accounts prior to
the early nineteenth century, it appears that woody forms of vegetation have
always been scarce on the island (Brown et al. 1989). Britton and Morton
(1989) point out that this is HOt typical of all barrier islands as those of the
rt oak- or pine- dominated woodlands.
Arid climate is a likely factor in the lack of woody vegetation but is not the
only factor as a number of northern Mexico barrier islands have dense stands
of thorn-scrub woodland (Britton & Morton 1989). Northern Padre Island
may have supported some scattered oak mottes that have declined through
time (Britton & Morton 1989; Brown 1989) but is primarily a grassland
ecosystem. Therefore, central and southern Texas barrier islands are unique
in having a herb-dominated flora and PINS is an excellent example of this
type of ecosystem. PINS also has the richest flora of the four barrier island
ecosystems that have been studied in Texas (Nelson, unpublished data). A
forthcoming investigation will use data from this checklist to elucidate a
common flora for four South Texas barrier islands, ity similarities
among the islands, and discuss unique floral components for Marizorda Island,
Mustang Island, northern Padre Island, and southern Padre Island (Nelson,
unpublished data).

eastern Gulfand Atlantic

CHECKLIST OF THE VASCULAR FLORA OF PADRE ISLAND NATIONAL SEASHORE

Plants that are woody are listed as such; plants without this designation
are herbaceous in habit. Habitat designations: coppice dunes (CD), foredunes
(FD), barrier flat (BF), and tidal flat (TF) (Nelson et al. 1997, 1999).

CANTHACEAE AIZOACEAE
Ruellia corzoi Tharp & Barkl., PINS, Lee 2532. Mollugo verticillata L., Indian chick weed.
AGAVACEAE ene 499,
ce constricta Buckley, Beargrass, Spa an- Sesaviam maritimum (Walt.) Britton, Sterns,
ish bayonet. PINS, Jones 756. Shrub. B & Poggenb., Sea purslane. NSA; Rechenthin
anc ED: and Passe
Yiuca trecitleana Cart., Spanish dagger, Palma Sesuvinm ee een (L.) L., Cenicilla.

es (1966), Rabalais (1975). TAIC, Nelson and Odvody N293.
en BF ie FD. Sesuvium trianthemoides Correll, Sea purslane

1232
TAC, Nelson, Macke, and Wilburn N697.
BE.
Sess. m Rat., Sea purslane. PINS,
is 6400 JO.
Tr. pei is ss wm L., Horse purslane.
S, Jone
ALISMATACEA

Echinodorus beteroi

—

Spreng.) Fassett, Burhead.
133.

aaa THACE _

_M. Johnst., Sandhills
amaranth, PINS, 6662
Amaranthus blitum L., ay Rabal lais (1975),
ry
CD

Amaranthus greggii S. Wats
Amaranth. TAIC, Ne/son - 5
Amaranthus ee ; a ama-
rth. PINS, S00.
Blutaparon vermicul oh Mears sakes
TAIC a elson and i asby N398. TE
Celosia re Vahl, PINS, ai 2470.
Snake cotton,
966).

Froelichia drummondit Mog.,
Cot veed. NSA;

Proelichie acai (Nutt. ) Mog. var.
floridana, Snake cotton, Cottonweed.
PINS, 6689.

Tidestromia lanuginosa Nutt.) Standl. var.

ones (19

lanuginosa, Espanta vaqueros. TAIC,
i417. BE

Nelson N 4 /

APIAC
Cente le astatica (L.) Urb., PINS, Joes 893.
Hydri cor \VlOC DONAVIEHS TS is Commers. ex Lam. F

Sombrerillo. TAI C, Nel and Perez-
Johnson N318. FD

Hydrocotyle ei. tes de Ve-
nus. PINS, Jones 1390.

ASCLEPIDACEAE

a ee emor es aie Vail, Milkweed,
Silkweed. NSA; Jones (1966).

Asclepias en Cham. & Schledl.,
M - ya de zizotes. TAIC, Nelson N328.

ie viridiflora Rat., Milkweed, Silk-
weed. NSA; Rechenchin and Passey
(1967)
. ici Pers., PINS, 924.
ASTERAC
Ambrosia arte ae Lb:
Altamisa, NSA; Bae us and Horton

Short ragweed,

Stipa 18(4)

(1979), Higginbotham (197 2), Rabalais
(197 bee

Naa), ps

, Western ragweed.
eg a peas N352. FD,
nd T

TAIC
BF,

Aad dracunculide (DC.) Nutt

sroomweed, Snakeweed. NSA, Ri edad als

(197

i sceee skirrhobasis (DC.) Trel. var.

s, Lazy daisy. TAIC
Nelson and Guerra a 375

Baccharis neglecta Britton, Roosevelt weed,
New Deal weed, Jara dulce. PINS, M.
1, 7227. Shrub.

Baccharis salicina Torr. & A. Gray, Seep

thalassius Shinne

willow, Jara, Water-wally.
Rechenthin and eae (1967). Shrub.
— es scens (L. ., Sea ox-eye daisy.
sie Nelson ue ‘idee N305. Sub-
shr a to shrub. BF and TF.
Basketflower,

.NSA

Centaurea americana Nutt.,
ae thistle, Cardo del valle
achin and Passey (1967).
Pea eee UtE., Golden aster, Cam-
Fee ieee - .C. Museum, Jones 6205.
Michx. var. e//ottir Torr.
& he City, ‘Bull thistle, Yellow thistle.
TAC, Nelson, Macke, and Wilburn N717.

BE.

Cirsium texanum Buckley, Thistle, Plumed
thistle. NSA; Rabalais (1975

Conoclinium betonie tfolinm (Mill) King &

Rob >, Nelson and Martinez N38.
FD, B E and TE.

Conyza canadensis (L.) Cronquist, Horse-
weed. PINS, 792

Conyza ramosissima Cronquist, NSA; McAtee

Coreopsis tinctoria Nutt., Coreopsis, Tick-
seed, ee Nelson, Macke, and Wilburn
N655

Croptilon diner iat Nutt. ) Raf., Scratch-
daisy. ag GS :

Crp rigiifolion B. Sm.) E.B.Sm.,

mn Macke, and

—
Ns
a

Scratch-daisy.

Wilburn NO3O. FIL
Dyssodia ea ‘ve ) Robins. var.

7 ntachaeta, Parralena. TAC, Ne/son.
Macke, and Wilburn x 648. BE.

)

NEGRETE ET AL.,

Dyssodia tenuiloba (DC.) Robins. es
oT ee ion TAIC, Perkins

Eclipta prostr LL CC. een , Jones
ca

— ie umbens (Houst. ex Mill.) Nesom,
Fle TAIC, Nelson N624. BF and

ED.
Exupatorium compositifolium Walter, Yankee
weed. PINS, Jones 5¢

ray eae (Torr. & A. Gray)
a) 64.

Pie verna ae ie t-tobacco, cael
PINS, Tie and Baker 04(

Plaveria browni? Powell, Longleaf i ria.
TAIC, Nelson N495. Endemic.
Gatllardia pulchella Foug., Indian blanket,
Ficcwheck TAIC, Nelson and Soto N361.

D, BF, and TE.

— falcata (Lam.) Cabrera, Cud-

verlasting. PINS, Jones and Baker

a
Gamochaeta pensilvanica (Willd.) Cabrera,
Cudweed, Everlasting. NSA; Jones (1966).
Helentum amarum (Raf.) H. Rock var.
an 2, C.C. Museum, ae 7927.
Helianthus #5 L., Common sunflower,
Mirasol. PINS, 548.
Helianthus argophyllus Torr. & A. Gray,
ilverleaf sunflower. PINS, 2516
Helianthus debilis Nutt subsp CHCHIN2EY, ifolius
Torr. & A. Gray) Heiser, Cucumberleaf
sunflower. TAC, Nelson, Macke, and
Wilburn NO84. TE

Helianthus praecox Engelm. & A. Gray subsp.
runyonii (Heiser) Heiser, Sunflower PINS,

2364.
“a sibaxillaris (Lam.) Bri
Rusby, Camphorweed. TAIC, Nelion and

es N307. CD and FD.

Iva annua L., Sumpweed, Marsh-elder,
Pelocote. NSA; Rechenthin and Passey
(1967).

lva ag itata Walte veed, Mars
NSA; es i ae es (197 Sf
Bat texensis R. R. Johnson, Sump-weed,
Marsh- a TAIC, Nelson N483. BEF

and TF.
Krigia occidentalis Nutt., Dwarf dandelion.
NSA; Jones (1966), Rabalais (1975).

Vascular plants of Padre Island National Seashore L2

Liatris elegans (Walter) Michx. var. elegans,
= feather, Se cee Blazing
tar. PINS, /
Fe eneinies an ephala (DC.) Shinners,
Camphor daisy. TAIC, Ne/son N420. BF
and TF.

Palafoxia hookeriana Torr. & A. Gray, PINS,
1383.

Palafoxia rosea (Bush) Cory var. rosea, NSA;

J 6).

Palafoxia texana DC. var. ambigua (Shinners)
B.L. Turner & M. I. Morris, C.C. Museum,
Jones 6770.

es es purascens (Sw.) DC., Canela. PINS,

ee peduncularis (Torr. & A. Gray)
Barnh. var. peduncularis, Mexican hat.
PINS, 530.
Rudbeckia hirta L. var. angustifolia (Moore)
erdue, Brown-eyed Susan, Cone flower.
NSA; Jones (1966), Rabalais (1975).
Senecio viddellii Torr. & A. Gray, Groundsel,
Ragwort, Squaw-weed. TAIC, Ne/son
902, FD
Solidago odora Aiton var. odorata, Sweet
goldenrod. PINS, /248

5 wee eae L., Seaside goldenrod.

PINS, Jones 125
Sonchus ee Acieoitadualce NSA;
(1966

Sonchus oleracens L., Sow thistle. oo Nelson,
Macke, and Wilburn N653. B
Symphyotrichum divaricatum dees : Nesom,
Hierba del marrano. PINS, Lee 2471.
Tetragonotheca repanda ae Small,
Rabalais (1975).
Thelesperma filifolinm ( ose ) A. Gray, Green-
thread. PINS, Whistler yale 1864.
piece nuecense B. L. Turner, Green-
thr PINS, Jones 1348,
Ver — wii (Cav.) Benth. & Hook.
ay, Cowpen daisy. C.C. Museum,

0 585.
Xanthinm strumarium L., Abrojo. NSA; Jones
(1966), Rabalais (1975).

AVICENNIACEAE

Avicennia germinans (L.) L., Black-mangrove,

Mangle blanco. PINS, 6668. Shrub.

1234
BATACEAE
ihe maritima 1., Vidrillos. TAIC, Ne/son
and Hornsby N393. Subshrub. TF.
BORAGINACEAE
Heltot um (Nutt.) A. Gray,
NSA; es ane Passey (1967).
Heliotropium curassavicum L. vat. curassavicum,

Seaside Vasa on ae co. is ae

Heli racemosum Rose & Meer
, He PINS, 5958.

—_—

iotrope.
BRASSICACEA
Cakile nena (Robins.) Millsp.,
, Laylor 398
Cakile lam ir (Willd.) Schulz subsp.
pseudoconstricta Rodman, ee Nelson,
Macke, and Wilburn N635.T
Lepidium lasiocarpum T. Nutt. ex con &
. Gray var. wrightit (A. Gray) C.L.
Pivehe. , Pepperwort, Peppergrass. PINS,

Sea

Jones 1378.

gee virginicum L., Lentejilla. TAC,
Macke, and Wilburn N652. BE.

tee nasturtium-aquaticum (L.) Hayek,

Water-c s. NSA; Baccus and Horton
(1979).

BUDDI EJACE Al

Polypremum procumbens L., C.C. Museum,

Jones 7568
CACTACEA

Opuntia ostmannt Salm-Dyck var.

cies (Enge B.D. Parfite &

Pinkava, Prickly pear, "Cholla . NSA; Jones

(1966), te 975), Rechenthin and
(1967). Shrub.

Opuntia humifusa (Raf.) Raf. var. bumifusa,
Eastern prickly pear. ah Jones (1966),
Rabalais (1975). Sh

Opuntia macrorhiza Enge ine var. macrorhiza,
Plains prickly pear. TAIC, Ne/son and
McKeller N355. Shrub. BF.

Opuntia stricta (Haw.) Haw., Prickly pear,
Cholla. NSA; Rabalais (1975). Shrub.

CAMPANULACEAE
Triodanis perfoliata (L.) Nieuwl., Venus
voking glass. NSA; Jones (1966),
Rabalais (1975).

Stipa 18(A)

CAPPARIDACEAE
Polanisia erosa (Nutt.) H.H. Iltis subsp.
breviglandulosa H.H. Itis, Clammy weed.
NSA; Jones (1966).
CARYOPHYLLACEAE
Silene antirrhina L., Sleepy catchfly. NSA;
Rabalais (1975).
sila oe a i Asch. &
aebn., Sand-spurr » Netson,
eh we a. NOS
Spergularia marina (L.) Griseb.,
sand-spurrey. C.C. Museum, Jones 6297.

Salt-marsh

CHENOPODIACEAE

Atriplex Maint ensis A,
centzo. NSA; Rabalais ao.

Atriplex pentandra Jacq.) Stand_., S altbush
NSA; Rabalais (197

Atriplex texana 8. See Saltbush. NSA;
Rabalais (1975).

Chenopodium album L., Lamb’s-quarters,
— NSA; Rechenthin and Passey

, Quelite

—

mibvostoides L. var. ambrostoides,
Mexican tea, Wormseed, Epazote. PI

2,

C penopediom Feel Mog., Pitseed

goosefoot. C.C. Museum, Jones 6298.

Sloman rr., Glasswort, oe
IC, Nelson and McKeller

“9

slo mta virginica L., Glasswor t aE
T Nelson rs French N367. TE.

Suaeda pap (Small) I. M. nes , Sea
blite, Seepweed. NSA; Rabalais, 1975
Suaeda linearis (Ell.) Moq., Sea Lice

5 S, 7139.
Suaeda tampicensis (Standl.) Stand|., Sea blite,

veed. NSA; Rabalais (1975).

Seepweed. PIN

CISTACEAE
Helianthemum georgianum oe Rock-
rose, Frostweed. PINS, °
Lechea mucronata Raf., - on PINS,
6628
CLUSIACEAE

Hypericum gentianoides (L.) Britton, Stems,
nb., Orange grass, Pineweed.
ee 2430.

Ue

~

-
U
-

th oO

Hypercinm hypericoides (L.) Crantz subsp.

NeGRETE ET AL., Vascular plants of Padre Island National Seashore P98)

hypericoides, St. Andrew’s cross. PINS,
Jones 721.

Hypericum pauciflorum Kunth, St. John’s wort.
PINS, O’Brien 2540.

COMMELINACEAE
C ; L. var. angustifolta (Mick hx.)

os , Widow’s-tears, day-flower. PINS,
52

ce erecta L. var. erecta, Hierba del
pollo. TAIC, Nelson and Hornsby N303.

an 4,
Tradescantia humilis Rose, Spiderwort,
spiderlily. PINS, 520.
CONVOLVULACEAE
Ipomoea — a Morning
glory. PI Lee 247
Ipomoea Sidon wah Gri seb., Beac
morning AIC, Nelson ne
Goodridge N289. = <_< FD.

Ipomoea ee Capra e (L.) R. Brown subsp.
brasil ne es Ooststr., aod vine,
Goa es ot morning glor A; Carls

al. (1991), ea praia
Kattner (1973), McAtee (1975), McAtee
and Drawe (1980), Rabalais (1975).
Ipomoea sagittata Poir., Morning glory. PINS,
ee 239]
CUCURBITACEAE
Citrullus lanatus (Thunb.) Matsum. & Nakai
var. citroides (L.H. Bailey) Mansf., Wa-
melon sandia. PINS, Jones 1352
Cvs var. melo, Muskmelon. NSA;
Jones (1966), ee (1975).
Cucurbita foetidissima Kunth, Buftalo-gourd,
illa loca. NSA; Rechenthin and
Passey (1967).
Ther villea lindheimeri (A. Gray) Greene,
y. NSA; Jones (1966).
a ian L. var. pendula, Meloncito.
seum, Jones 7140.
CUSCUTACEAE
Cuscitta oc Choisy, Pretty dodder. PINS,
Jones 885.

Cases es Engelm., Dodder, Love-
NSA; Rabalais (1975).
CYMODOCEACEAE
Cymodocea fuliformis Kutz.) Correll, Manatee-
grass. NSA; Rabalais (1975)

Halodile beandettet (Aen Hartog) den Hartog,

CYPERACEAE
teres Vahl, Flatsedge. NSA; Jones
(1966), Rabalais (1975).
Cy see echinatus (L.) A.W. Wood, Flatsedge.
NSA; Jones (1966), Rabalais (1975).
Cyperus eames L., Yellow nut-grass. TAIC,
Nelson N494. TF.

Cyperus haspan L., Flatsedge. PINS, Jones 1381.

Cyperus polystachyos Rottb., Flatsedge. PINS,
Whistler and Lee 241

Cy, _ pdtv (tenth, )R. Carter

Flatsedg ge. NSA; Jones (1966).

ce sino a var. a

m, Jones

on. retrorsus ts Chape, ade TAIC,
Nelson ;

Cyperus sae es — Talill o. NSA;
Carls et al. (1987), Carls et al. (1991).

Cyperus squarrosus L. var. squarrosus, Flatsedge.
NSA; Jones (1966), Rabalais (1975),
Rechenthin and Passey (1967).

Cyperus surinamensis Rottb., Flatsedge. TAC,
Nelson, Macke, and Wilburn N641. FD
and BF.

Eleocharis albida Torr., Spikerush. PINS,

Eleocharis arabe (Retz.) J. & C. Presl,

Spikerush. PINS, Jones 7410.
Eleocharts oe Torr., Spikerush. NSA;
Rabalais (1975)

Eleacharis flavescen (Poir.) Urb. var. flavescens,
o elson, Macke, and

tlh
Eleocharis geniculata - )J. J. Roem. & Schult.,
Spikerush. a Nelson, Macke, and
Elea aris interstincta oe els & Schult.,
rush. PINS, Lee 2475.
oo minima anth var, minima,
Spikerush. C.C. Museum, Jones 7417.
Eleocharis montevidensis &. Kunth, Spikerush
TA son, N348

Eleocharis parvila (Roem. & Schult.) Link
es & Schauer var. anachaeta
(Torr.) Svenson, Spikerush PINS, Jones 923
Fimbristylis cavoliniana (Lam.) Fern., TAIC,
Nelson N418. BE.

Pon elon iy
Nelson N419. BF.

Futrena es Michx.
Fall 7

sa ing Nan Umbrella-grass. PINS,

VICH 2

—

Michx.) Vahl, TAIC,

,C.C. Museum,

1D Paani MIC ane (Vahl.) G.C. Tucker,
P oe iG 89]

Rhy 16 ae ee Pfeitf., White-
aes aie grass. TAC, Ne/son,
Macke, and Wilburn Noe. BE

pee ae erectus Spas Palla os} Raynal,

sullrush. a it

Pree ie sonia . Pall

longespicatus (Britton) : . Sm., Sword-

a Var.

grass, Three-squared-bullrush. TAC,
Nelson ee alas Wilburn NO54. BF.
‘(Fern.) J. Raynal,
Bully NSA: Jo 966); Rechenthin
and souks
DROSERACE
Drosera bret a pa Sundew. PINS, Jones

ELATINACEAE
latine triandra Schkuhr. eeeaiiasee CC,
aes 6833.
EUPHORBIACE
se ge edn Town: Three-seeded mer-
cury. PINS, Jones 6560
oe aa texans (Mull. Aes Small, Bull
e, Mala mujer. C.C. Museum, Jones
9

nece
755

Cian yr “
Jones O54 1.

Croton fara Michx. var. /indheimeri
(Engelm. & A. Gray) Mill. Arg., ee
croton, Seca TAIC, Nelson, N42
BE.

roton ee ie roizat, PINS, Jones 959,

SL. war. lindheimeri Mii
Arg. " TAIC ‘ eae:

Croton endl asus L, var. prbentissimus Croizat,
PINS, 414.

y Michx., C.C. Museum,

ps

l,

Lee 2

Croton glandulos var. a Mill.
Arg., PINS, Hise 3 238

Crot eynus Croton. Michx.,
ie 1258.

Croton parksii Croizat, Croton. C.C. Mu-
seum, Jones 0670.

PINS,

Sipa 18(4)

Croton ae Jacq., Hierba del jabali,
Be TAIC; Nelson and Guerra,
N2 185. Subshrub. CD and FD.

Euphorbia al Jacq., Spurge. PINS,

Baker

ti io liott, Spurge. PINS,

ae bares L., Spurge. NSA; Carls
et al. (1987)
Euphorbia innocua Wheeler, Spurge. NSA;
s (1966), Rabalais (1975).
Euphorbia maculata L., Spurge. PINS, Jones
287.
Euphorbia serpens Kunth, Hierba de la
golondrina. C.C. Museum, Jones 7687.
Phyllanthus tll Be oe
Leat-Hower. PINS, Fa
Phyllanthus amides ex Spreny..
Leaf-flower. PINS, Jones and Baker 604.
Phyllanthus pudens Wheeler, Leaf-flower.
SA; Rabalais (1975),
ee sylvatica Garden ex L. subsp.
Hit, Q een sdelight PINS, Jones G17.
FABACEAI
Acacia minuata (M. E. Jones) Beauch. subsp.
munnata, Wuisache. NSA; Jones (1966),
Rabalais (1975), Rechenthin and Passey
(1967). Shrub or tree.
Aeschynomene oe M ; - X.,
Astragalus . oensis Dein
Loco weed. PINS, Jones 67
iting leptocarpus . ay, Milk-
etch, Loco weec S, Jones 7620.
Basi a Nutt. var. ance )
ndhi, Plains wild indigo.
rate (1975).
ee bracteata Muhl. ex Elliott var
laevicanlis (A. Gray ex Canby) Isley, Plains
wild indigo. TAIC, Nelson and Negrete
N402. BE
Canavalia rosea (Sw.) DC.,
Swordbean. PINS, Jones 91

var. abnormis,
250

Sticky
jointvetch.

Mil vetch,

ee

Cent) OSCMA UNI IAN) 1] (L. ) aan Bawenly
pea. PINS, Jones 747
Chamaecrista jst (Michx) Greene,

ee TAIC, Nelson N489. FD
and BE

Clitoria ae L. PINS, Lee 2400.

NEGRETE ET AL.,

Crotolaria sagittalis L., Rattlepod. PINS,
Jones 4092

Datlea emarginata (Torr.& A. Gray) Shinners,
PINS, Nelson, Macke, and Wilburn N7 13.

Daly lanata Spreng., Woolly dalea. PINS,
Jones and Baker 6562.
Datlea obovata (Torr. & A. Gray) Shinners,
Pussyfoot. PINS, Jones 738.
Dalea pogonathera A. Gray, Hierba del
corazon, Bearded dalea. NSA; Carls et
1987

Desmanthus virgatus (L.) Willd. var. depressus
Humb. & Bonpl. ex Willd.) B.L. Turner,
PINS a ue 850.

Eryth rbacea L., Coral bean, Colorin.

__ PINS, tah

s Benth., Hoary milkpea.
PINS, La 2 420.
G ee tia marginalis Benth., PINS, Whistler
524.

Gloom vesicaria (Jacq.) R.M. Harper,
eum, Jones 7748
Indios miniata Ortega vat. elebionebals
.ex Torr. & A. Gray) B.L. Turner,
seals pea. NSA; Drawe etal. (1981),
inbothan (1972), Jones (1966).
/ ce nuniata Ortega var. miniata, Scarlet
pea. TAIC, Nelson and Hornsby N301.
D and BE
Indigofera suffruticosa Mill., C.C. Museum,
Jones 6673. Shrub.
Leucaena leucocephala (Lam.) de Wit, C.C.
um, Jones 7924. Shrub or tree.
Medicago polymorpha L., Bur-clover. PINS,
Jones 7
Melilotus albus Medik., White sweet clo-
ver, Huban. PINS, ae and Lee 248,
Melzlotus ies (LAE. ., Sour clover,
Alfalfilla. TAC, ee Macke, and
Wilburn N692. BF.
Mimosa latidens ee ll) B.L. Turner, Sen-
sitive brie NS, Whistler 1837
Mimosa str a Torr. & A. Cu
ne lerpuff, Vergonzosa. NSA; pee
1966), Rabalais (1975).
Neptunia lutea (Leavenw.) Benth., Yellow-
A; Rechenthin and Pisey (1967).
Neniunia pubescens Benth. var. pubescens, PINS,

Vascular plants of Padre Island National Seashore Lea

Jones and Baker 741.
Parkinsonia aculeata L., Retama. C.C.

useum, 79D096. Tree.
Pediomelum rhombifolinm (Torr. & A. Gray)
C.C. Museum, 3563.
Prosopis glandulosa Torr. var. glandulosa,
Honey mesquite. PINS, Jones 9/2. Shrub
or tree

Rhynchosta americana (Houst. ex Mill.) Metz,
Snoutbean. TAIC, ae and Goodridge
N321. FD and B
Rhynchosia minima in DC., Snoutbean.
TAIC, Nelson and Perez—Johnson N306.
bet and a
Rhynck is 1) DCE., Dollar-
ie NSA: Reshentinaed Passel 967).
Rhynchosia senna Gillies ex Hook. var. texana,
Snoutbean ee C, Nelson, Macke, and
Wilburn N704. BE.
Seshania dr ae (Rydb.) Cory,
Rattlebush, Poison bean, Coffee bean.
PINS, Jones 914. Shrub.
Sesbania exaltata (Raf.) Rydb. ex A. Hill,
Bequilla. PINS, Tarp 233. Subshrub.
a a var. occidentalis (L.) Isely.
hora. PINS, Tharp 40. Shrub
Str Pie held ula (L.) Elliot, Amberique

ode
Strophostyles leiosperma (Torr. & A. 7 oo
Sleek-seed bean. PINS, Jones 9
Stylosanthes viscosa Sw., Pencil- he | NS,
Lee 2385.
Vigna luteola Jacq.) Benth., PINS, Jones 750.
Zornia bracteata J. F. Gmel., Viperina. PINS,
Thistler and Lee 2401

FAGACEAE

Quercus minima (Sarg.) Small., C.C. Mu-
m, Jones 7922. Shrub.
Quercus virginiana Mill. var. fuszformis (Small)
ou TAIC, Nelson N562. Shrub
or tree. BE.
ee virginiana Mill. var. cies Live

o. PINS, Jones 937. Tre

oak, enc

FUMARIA

EAE
a . Gray)
ray s eee texensis G. Ownbey,
Scrambled eggs. NSA; cane (1966),

Rabalais hos,

1238
GENTIANACEAE
eee exaltatum (1...) Salisb. ex G. Don,
chfly-gencian. PINS, Jones 883.
§ at arenicola Greenm
C, Nelson N625. FD a
Sari campestris Nutt.,
, Nelson N5O9. TE.

, Rose- oe
nd B
cee -gentian.

GOODENIAC

E
Scaevola ae (L.) Vahl, PINS, Tasner

1834. Subshrub.

HY DROCHARITACEAE
Halophila engelmannii Asch., PINS, Lee 2448.
Thalassia testudinum Banks & Soland. ex
Konig, Palmas del mar, Turtle-grass.
NSA; Rabalais (1975)
HY DROPHYLLAC
eka elie a NSA; Jones (1966),
Rabalat
share vatuliflora (aed: & A. Gray)
Gray, NSA; Rabalais (1975).
IRIDACEAE
Sisyrinchium biforme E.P. Bicknell, Blue-
eyed grass. TAIC, Nelson and ee

—

N3806. BE
Sisyrinchinm oe eee P. Bicknell, Blue-
eyed grass. PINS, Jones 724.

JUNCACEAE

s Michx., Rush, Bog-rush.
PINS, ca 818.

ee a? Engelm.,

. NSA; Jones (1966).

tes Ws sin L. var. ee Toac

Ce. seu, Jones 18 Se

Rush, Bog-

LS
jon

-rush.

4
Sc

Juncus mar. ae Rostk., Rush, Bog-rush.
PINS, ae C, Nelson, es and Wilburn

N664
Juncus eee M. A. Curtis, Rush, Bog-

rush. 3 Nelson, Macke, and Wilburn
N699.

Juncus roemerianus Scheele, PINS, Whistler
2484

Juncus scspoiees Lam., Neddlepod rush
TAIC, Nelson N513. BE

Juncus validus Coville var. fascinatus M.C.

st., Rush, PINS, Jones

6864.

|
=)

Bog-rush.

areas AE
/ tata L. var. lasiodonta A. Gray,

Stipa 18(4

~~

Spotted beebalm, Horsemint. C.C. Mu-
seum, Jones 7099.
Scutellaria drummondii Benth., Skullcap.
Jones (1966), R Rabalins (1975).
Scutellaria muriculata Epl., Skullcap. PENS,
ones O56
eee: var. cubense, Germander.
SA; Rechenthin and Passey (1967).
LAURACEAE
Persea borbonia (L.) K. Spreng., Red bay.
PINS, Jones 905. Shrub or tree.
LENTIBULARIACEAE
Utricularia subulata L., Bladderwort. C.C.
Museum, Jones 06827
sip
Cooper: nmmondit Herb., Cebolleta. NSA;
Raval ais (1975).
Nothoscordum bivalve (L.) Britton, Crow-
1. NSA; Jones (1966).

LINACEA

ee (Small) H.K.A. Winkl., Flax.
NSA; Drawe et al. (1981), Jones (1966),
Rabalais(1975).

Linum lind Engelm.,
Rabalai

Linum ns (Raf.) Shinners, Tufted

. NSA; Rabalais (1975).

Flax. NSA;

LOASACEAE
Mentzelia oligosperma Nutt. ex Sims, Stick-
leaf, Chicken thief, Pegajosa. NSA; Jones
(1966), Rabalais (1975)
LYTHRACEAE
Ammannia latifolia L., C.C. Museum, Jones
pe 7
Lythrum adlatum Pursh var. lanceolataum
iott) Torr. & a
Loosestrife. TAIC, Nelson N424. TE.
Ly Hit aca hile aot & A. Gray, Hierba
del , Whistler 2469.
Ruale ramosior ; ‘ ‘Bochue.
S, Jones 08351.
MALVACEAE
po roa ed (L.) K. Presl ex A. Gray,
Salt marsh-mallow. C.C. Museum, Jones
6912.
Matlvastrum americanum (L.) Tort.,
loca. PINS, Jones 916.

—
ies!

Tooth-cup.

Malva

NEeGRETE ET AL., Vascular plants of Padre Island National Seashore

Matvastrum coromandelianum (L.) Garcke,
S, Jones 917
Sida ciliaris L., Bracted sida. Whistler and
Lee 2417

Sida cordifolia L., NSA; Rabalais (1975).
Sida lindheimeri Engelm. & A. Gray, PINS,
Jones 868.

Sphaer a lindbeimeri A. Gray, False mallow,
obe

PINS, Baker 763

mallow.
MYR
fae ae (L.) Small, Wax-myrtle,
Candle-berry. PINS, Jones 764. Shrub.
NAJADACEAE
Nayas sosieeuc dala pe ) Magnus,
C

er-nymph. PINS, Hamby
pe ene 1152.
NYCTAGINACEAE
Boe niet coccinea Mill., Scarlet spiderling.
ane - 5).

Mivabilts er) Heimerl., White
ari eee Gf 7e.
ONAGRACEAE
se ei berlandieri Spach subsp. pinifolius
gelm. ex A. Gray) Towner, NSA; Jones
aoe.
Cal i aw serrulatus (Nutt.) P.H. Raven,
ening primrose. ne Nelson
and Perez-Johnson N343.
Gaura longiflora Spach, NSA; ce (1966),
Rabalais (1975).
Gaura sale iflora Douglas ex Lehm., Liza
vet-leaf gaura. NSA; Jones ( oe
saris (1975)
Ludwi igia glandulosa Walter, Cylindric-fruited
ludwigia. NSA; Jones (1966), Rabalais
(1975):

Ocenothera drummondit Hook. subsp. drum-
mondit, ing primrose. TAIC,
Nelson and Perez- Pte N287. CD and
FD

coe een Smyth, Evening
primrose. NSA; Jones (1966), Rabalais
(1975

Ocnothera laciniata Hill, Cut-leaved evening
primrose. TAIC; Ne/son N566. BF and
TE

Ocnothera speciosa Nutt., PINS, Hambly 1195.

[a0

ORCHIDACEAE
Spivanthes vernalis Engelm. & A. Gray, Spring
adies’ tresses. TAIC, Nelson and Soto
N329. FD and BF
OROBANCHACEAE
Orobanche ludoviciana Nutt subsp multiflora

tt.) Collins, Broomrape. C.C. Ma =
seum, ae 2040.
OXALIDA
Ol dili o subsp. di/lenii, Wood-
rrel, lady’s sorrel. NSA; Jones (1966),
Rabalais (1 O75
Oxatis drummondii A. Gray, Wood-sorrel,
rel. NSA; Rabalais (1975).
PAPAVERACEAE
Argemone albiflora Hornem. subsp. texana
wnbey, White prickly poppy.
NSA; Higginbotham (1972).
Argemone sanguinea Greene, Red poppy.
NSA; Jones (1966), Rabalais (1975).
PHYTOLACCACEAE
Phytolacca americana \.. vat. americana,
Pokeweed, pokeberry. C.C. Museum,
Jones 7657.

PLANTAGINACEAE
Plantago oo Nutt., Plantain. PINS,
Jones 68C
ae ae Fisch. & C.A. Mey.,
Tallow weed. A Macke, and
Pe ciedss N 6

ie oo plantain.

TAIC, Ricon eee

PLUMBAGINACEAE
Limonium carolinianum (Walter) Britton,
Sea-lavender, Marsh-rosemary. TAIC,

Nelson N518. TE.

POACEAE
Andropogon gerardii Vitman, Big bluestem.
INS,
Andropogon glomeratus (Walter) Britton,
ey & Poggenb., Bushy ee
Nelson and Recadee N381. TE

Ps ternarius Michaux vat. ternarius,
Splitbeard bluestem. PINS, Jones 1312.
Andropogon virginicus L., Bieopte ae PINS,

Jones 903

1240

Aristida longespica Poir. var. geniculata (Raf.)
Fern., Three- awn grass. TAIC, Nelson
N499.,

Ariens Mc, Prairie three-awn.

Rechenthin and Passey (1967).

Aristida yee Nutt. var. parpirea, Purple
three-awn. NSA; Rechenthin and Passey
(1967).

Arundo donax L., Giant reed, Carrizo,

seorgia cane
Wilburn NO32. BE

Avena fatua L., Oats. NSA; Jones (1966),
Rabalais (1975).

Bothriochloa barbinodis (Lag.) Herter var.

j TAIC, Guerra

barbinodis, Beardgrass.

OOG06.
Bothria ee L. var. songarica (Fisch.
and € .) Celarier & Harlan, PINS,

ieee op 2,

Bothrtochloa lagurotdes (DC.) Herter subsp.
ieee (Steud.) Hack., Silver beardgrass,

er ius PINS, Jones 7684.

Br tl Vahl, Rescue grass. NSA;
_ Jone Oo) eo ais (1975).

, Cadillo. TAC, Nelson,
Macke, and W es N7006. FD and BF.

Cenchrus spinifex Cav., Grassbur, Coast
sandbur. TAIC, Nedson and Odvody N3 15.
3 BE.

ane

/ropogonoides Fourn., Fingergrass.
PINS, ae 6732.
Chloris canterai Arech. vat. canterai, Finger-
A; Jones (1966), Rabalais (1975).
Ci CHE a ita Bisch., Hooded fingergrass,
Hooded windmill grass. NSA; Jones
(1966), Rabalais (1 a ares and
Passey (1967).
Chloris gayana Kunth, R
Jones 7147
Chloris vertic ae Nutt., Windmill finger-
grass, Tumble windmill grass. NSA;
Rechenthin and Passey (1967).
Cynodon dactylon (L.) Pers. var. dactylon,

hodes grass. PINS,

Bermudagrass, Pata de gallo. PINS, Fa//
242
Dactylocteninm aegyptinm (L.) Beauy., Crow-

foot. NSA; ms 966), Re aie
Dichanthinm annilatum (Forssk.) Stapt var.
annulatum, PINS, Whistler 2480.

TAC. Nelson, Macke, and

Stipa 18(4)

Dichanthiiwan

(Poir.) C.E. Hubb.,

C. Museum, Jones 6733.

Digitaria arenicola (Swall.) A. Beetle, Crab-
grass. TAC, Nelson, Macke, and Wilburn
N713. FD.

a

Digitarh 4 bicornis (Lam.) Roem. & Schult.,
C useum, Jones 7902
Digitaria cognata (Schult.) Bits g., Crabgrass.
PINS, Fall 1253.
Digitaria sanguinalis (L.) Scop., Northern
crabgrass. NSA; Rechenthin and Passey
1967)

wis He de rexana Hitche., Crabgrass. PINS,
‘a

Distichl is eat ) Greene, Saltgrass. PINS,
Whistler

Et ee crus-galli (..) Beauv. var. cris-
valli, Barnyard grass. NSA; Rechenthin
and Passey (1967).

Echinochloa crus-pavonis (Kunth) Schult.
NS echenthin and Passey (1 967).
Echinochloa walteri (Pursh.) A. Heller, NSA;

_ Rec eee and Passey (1967).
lier? Daveau, Lovegrass. C.C.

M steed 7582.
Eragrostis capillaris (L.) Nees.,
_NS SA; Ri anions ‘ mance
fi / Buckley, Gummy
love grass. NSA; Rechenthin and Passey
(1967).
Eragrostis intermedia Hitche. var, stermedia,
es rass. NSA; Rechenthin and Passey
967).
Paes /ngens Nees., Lovegrass. TAIC,
Estrello

Lacegrass.

Eragrostis pectinacea (Michx.) Nees.,
Lovegrass. NSA; Rechenthin and Passey
(1967)
E ae wc J. Presl pane a
(Torr.) S.1 1, Red lovegra
Nelson, an Wilburn ae Cc -
FD, and BE.
Eragrostis sesstlispica Buckley, Lovegrass.
NSA; Jones (1966), Rabalais (1975),
schenthin and Passey (1967).
E pikes spectabilis (Pursh) Steud., Lovegrass.
NS, Jones 1586
E Se s petrea (Sw, ) Desv., TAC, Ne/sov,
Macke, and Wilburn N642. BF.

a

NEGRETE ET AL.,

Hordeum pusillum Nutt., Little barley. TAIC,
3) CE,

Leersia hexandra Sw., PINS, ee

ee dubia (Kunth) Nees., Green

sprangletop. NSA; Jones C1966) Rabalais

Leptochloa fascicularis (Lam.) A. Gray var.
fasicularis, Sprangletop. NSA; Baccus
and Horton (1979), Jones (1966),

Rabalais (1975).

ee mucronata (Michx) Kunth, Red
sp etop. ; Rechenthin and
Passey (1967).

Lolium perenne L., Ryegrass. NSA; Jones
(196 Rabe (1975).

Monaashooblee littoralis Engelm., TAC,
Ne Macke, and Wilburn N67, TE.

aes Cel (Lam.) Trin., Gulf

hly. PINS, Wéistler 2502.
Nassella leucotricha (Trin. & Rupr.) R. Pohl,
exas speargrass, Texas winter-grass.
NSA; Jones o od

Panicum acicilar x Poir. var. aneusti-

lium (Elliot) ei S. a Jones, PINS,
ones 702

Panicum eat Sw., Panic grass. PINS,
Jones

amarum Elliot var. ae (A.
ae & Chase) P. Palmer, Beach panic.
TAC, Nelson N624. CD nd FD

Panicum amarum Elliot var. amarum, Beach

a AIC, Nelson N493.CD and FD.
Pani ale ude Vasey. Panic grass.
Nelson and ee N309, FD,

es nebo Hitche. & Chase, Panic

S, Fall

grass 56 1.
Panicum Uieanlbes Schult., PINS, Whzstler
3.

ee portoricense Desy. ex W. Hamilt.,
Panic grass. TAIC, Ne/son and French
N369. BE.

Panicum scare Elliott, Panic grass.
es elson N560. BE.

PINS, Whistler 2508.

Ppa a ) C.E. Hubb., Sickle-

, Alaniz.

Pa. iat petostcalesun Vasey, Paspalum.
AIC, Nelson, N486. FD and BF.

Vascular plants of Padre Island National Seashore 1241

Paspalum plicatulum Michx. var. plicatum,
Brownseed paspalum. PINS, Jones 899.

Paspalum setaceum Michx., Paspalum. TAIC,
Nelson and McKeller N299. FD and BE.

Paspalum vaginatum Sw., Paspalum. PINS,
Jones 18%

Pennisetum ciliare (L.) Link var. ciliare,
Buffelgrass. TAIC, Ne/son and Hornsby,
N297. FD.

Sn om glaucum (L.) R. Br., NSA;
Rechenthin and Passey (1967).

Phragmites australis (Cav.) Trin. ex Steud.

wn
co
S

sp. australis, oe feed, GC.
Muscun, Janes O76

P, oo ~, Rabbitfoot
ass. TAC, Neon, eee i Wilburn

NO6S80. TE.
SUennada haar (Nutt.) W. Trel.,
Tumblegrass. NSA; Jones (1966),
Rabelais Cee
(Michx.) Nash var.
lint (Nash) Gould, Seacoast bluestem.

AIC, Nelson and Cadena N314.FD an

rd,

Setaria leucopila & Merr.) K. Schum.,
NSA; Jones (1966), Rabalais (1975).

Setaria macrostachya Kunth. NSA;
Rechenthin and Passey (1967).

Setaria parviflora (Poit.) Kerguélen, NSA;
Jones (1966), Rabalais (1975),
Roenthip we Passey (1967).

So (L.) Nash, PINS, Whistler

— halepense (L.) R. Br., Johnson grass.
ce Jones 8) Rabalais (1975).
Spar Lois. var. glabra (Bigel.)

C.C. Museum,

ern.,S i
Jones 6900.
S, a ees (Ait.) Muhl.,
ordgrass. TAIC, Ne/son N42
eae ert (Trin.) Merr. ex ie,
Sacahuista, Gulf cordgrass. C.C.

ais eadow

seu “es 4718

Sr iepholie o Dare (Michx) EF. Lamson.,
Prairie wedgescale. NSA; Jones (1966),
Rabalais (1975)

Sporobolus airoides Torr.) Torr. subsp. airoides,

Alkali sacaton. NSA; Jones (1966),

1242

Rabalais (1975), Rechenthin and Passey
a)
Sporobolus ees (Torr.) A. Gray, Sand
dropseed. ; Carls et al. (1987), Jones
(| = cere (1975), Rechenthin and
Passey (1967).
Spor sholus pyramidatus (Lam.) Hitchc.,
Dropseed. PINS, P
Soros hati Hitche., Coastal sacaton.
IC, Nelson NO88. Endemic. CD and

FD.
Sporobolus virginicus (L.) Kunth, Coastal
sey ae TAIC, Nelson and Hornsby
N3 FD, BE, and TE
Tio se N Foie,
Rabalais (1975).
rr aii Duis Sil
PINS, Jones 6719.
a (L. H. re a Tridens.
Nelson NSIS.
wee a aa (Walker) Chapm. var.

, NSA;

Jones

, Silveus-grass.

purpurea, P urple sandgrass, PINS, Jones

S896.
Uniola Span L., Sea oats, Espiga del
mar. TAIC, Nelvon and Cadena N291,

CD and =

Urochloa ere (Buckle .D.
Webster, TAC, Ne/son, Macke, and W ae
ae FD

Urochloa texana (Buckley) R.D. Webster,
PINS, Whistler 2507.

Vaseyochloa multinervosa (Vasey) Hitchc.,
Texas grass. PINS, Jones 6713.

Vulpia octoflora (Walter) Rydb.,
fescue. TAIC, Kiddy.

Six-weeks

POLEMONIACEAE
Phlox drummondii Hook., Ph
Nelson, Macke, and Wilburn 7 A
Phlox glabriflora (Brand) W
ora, Rio Conde phlox. C u-

W hitchouse subsp.
glabriflo C.M

Yo

seum, Jones 07
Phlox glabriflora a Whitehouse subsp.

/ittoralis (Cory) hg Rio Grande
PINS, Le

phlox,
POLYGALAC
Poly fe we <a"
T , Nelson NO26. FD and BF
a imarnata L., Polygala, Milkwort. TAC,
Nelson, Macke, and Wilburn NO37. BE.

Polygala, Milkwort.

Sipa 18(4)
Polygala verticillata L., Polygala, Milkwort.
C.C. Museum, Jones 1980.

— GON on io
oli]

2 Nutt. var. BG
Wild ee NSA; McAtee (1975),
Rabalais (1975).
B se a multiflorum Benth., Wild buck-
wheat. TAC, Nelson, Macke, and Wilburn
oo FD al BE
Polygonella polygama (Vent.) Engelm. & A.
Gray, October-flower. PINS, Jones 6698.
ers ae EAE
al subsp oleraced, —
Series. PINS. Jones and Baker 78
Portulaca pilosa L.., C

Jones 2339-A.

a"

usme. C.C. <tie

Talinum se ag Nutt., Flame-flower.
. Museum, Jones 2243.
PRIMULACEAE
Anagallis arvensis L., Pimpernel. TAC,

Nelson, Macke, and Wilburn NOS1. BE.
C.C.

— minima L., Chatfweed.
Museum, Jones 0805

ca Pee ae Water-pimpernel,
B

>, Nelson and Odvody

ee

N350. BF and cc
RUBIACEAE
oe occidentalis L.
en »mmon buctonbush, Honey-balls,

var. californicus

Pau sie . PINS, Jones 908. Shrub
or small tree

Diodia teres Walter var. teres, Poor Joe, Rough
buttonweed. PINS, Jones 6559,

— otis nigricans (Lam.) Fosberg, Bluets. a

SON, es and Wilburn NO85
>. Wright ex A. ee
TAIC, Nelson and

A. ‘Gay, Blues.
Martinez N335.
Oldentandia boseit ee ) Chapm., Bluets.

892
Oldenlandia panes L., PINS, Jones 7441.
Richardia brasiliensis Gomes, PINS, Jones

RUPPIACEAE
Ruppia maritima ., Widgeon-grass. PINS,
Jones and Baker 6552.
RUTACEAL
Zanthoxylum clava-herculis L., Pepperbark,

NEGRETE ET AL., Vascular plants of Padre Island National Seashore

Hercules-club, Prickly ash, Tickle-tongue,
Toothache tree. PINS, Jones 7058. Shrub
or sma
ee hirsutum Buckley, Tickle-
oothache tree, Prickly ash. PINS,
Jones 786. Shrub or small tree.
SALICACEAE
Salix nigra Marshall, Black os Satz.
AIC, Nelson N563. Tree
SAXIFRAGACEAE
Lepuropetalon spathulatum Elliott, C.C.
useum, Jones 0808.
SCROPHULARIACEAE
Agalinis fasiculata ae Raf.,
Museum, Jones 75
A aie heterophylla pe ae ex oF
Molby

CC.

rie agalinis. TAIC, B

oe

Agalinis maritima (Raf.) Raf. var. glandiflora
(Benth.) Shinners, Seaside gerardia, Salt
marsh gerardia. TAIC, Ne/son and Negrete
N411. BE

Agatinis strictifolia (Benth.) Pennell, PINS,
Lee and Whistler 2390

Bacopa caroliniana (Waleer ) Robins., Blue
hyssop. b ; Rechenthin and Passey
(1967)

— monniert (L.) Pennell, Ree

PINS, Jones and Hambly 1157.

Bacopa rotundifolia (Michx. oa PINS,
Whistler 2467.

‘icana L., Bluehearts. TAIC,

Nelson N623. BE.

uchwerd aner

Castilleja indivisa Engelm., Indian paintbrush,
Paintedcup. NSA; Rechenthin and Passey
(1967).

nia dubia (L.) Pennell var. anagallidea
.) Cooperr., False pimpernel. PIN
Jon 5506.
rbi mb. & Bonpl.
ex Willd. ‘bsp pe ey Snap-
e. PINS, ans ae me 790.

+}

dragon v

a ee texans (
Texas toad- aan NSA; cae aocee
Rabalais (197

tton

>

Stemodia lanata & Moc. ex Benth
Woolly stemodia. TAIC, Ne/son dea
Negrete N409. FD and BE.

1243
seri naan
cum annuum L. var. aviculare (Dierb.)
D A fo pat ce Eat | 1
guin, Cals
(1975). Sie or shrub.
Lycium carolinianum Walter var. quadr
(Du oa Hitchc., Carolina wo . rry.
C, Nelson ee0. Shrub. T
Grah., Tree —

Bird pepper. Chilipe-
Rabalais

we glauca
Mustard tree, Rapé, Gigante, Buena
moza. C.C. Museum, Jones 7584. Shrub
or small tree.

Physalis cinerascens (Dunal) Hitche. var.
spathulifolia (Torr.) J. Sullivan, Ground
cherry. TAIC, non and Martinez N290.
oD: FD, and B

So, Mill., American night-

shade, Hierba mora negra. PINS, Jones
309.

3
Solanum rostratum Dunal, Buffalo bur,
Kansas-thistle, Mala mujer. NSA;
Rechenthin and Passey (1967).
TAMARICACEAE
Tamarix aphylla (L.) Karst., Tamarisk, Salt
r. NSA; a heii Tree.
ee canartensis W narisk, §
A; Rabalais fe iris or tree.

, Salt cedar.

Weg abe 7s Lour., Tamarisk
C.C.M

useum, Jones 8147. Shrub or smalll

tree.
Tamarix gallica L.., Tamarisco, Rompevientos.
Jones 0551. Shrub or small tree
Tamarix ramosissima Ledeb., Tamarisk, Salt
PINS, Whistler and Lee 2395. Shrub
or small tree

TYPHACEAE

oe Pers., Tule. PINS, WAistler
and Lee 243
Typha ee i , Common cat-tail, Tule
espac INGA: Higginbotl vam (1 972),
scat hin and Passey (1967)
URTICACEAE ~
ria fl t., Pellitory. NSA;
McAcee io es Rabati (1975).

‘@ Muhl. ex Willd. var.
obtusa ee rdb. ex Small) Shinners,
Hammerwort. PINS, Jones 795.

Urtica chamaedryoides Pursh, Ortiguilla.
NSA; Jones (1966), Rabalais (1975).

1244

VERBENACE
Glandularia naiti (Nutt.) Nutt. var.
bipinnatifida, PINS, Jones 1205
Lantana camara L. Indian anton,
. Sabla bedions: NSA; Rabalais (1975
966). 7
ee urticoides Hayek, Texas lantana,
Hierba de cristo, Calico bush. PINS, Joves
7310. Shrub.
Phyla lanceolata
frog-fruit
(1 967 By:
Phyla nodiflora (..) Greene, Common frog

Xechenchin and Passey (1

Michx.) Greene, Northe
SA; Rechenthin and ee

Zit

fruit, Cape-weed, Turkey-tangle, Mat-grass,
Hierba de la Virgen Marta. TAIC, Nelson
and Soto N320. FD and BE.

Sipa 18(4)
Verbena halei Small, Texas vervain. PINS,
Jones 7 }
VITACEAI
Cassius incisa Des Moul., Marine-ivy treebine,
Cow-itch, Hierba del buey. PINS, Joes

——

771
XYRIDACEA
Nyris jupicat Rich, Yellow-eyed grass. PINS,
Jones 7396
ZYGOPHYLLACEAE
Kallstroemia hirsutissina Vail ex Small,
varpetweed, C.C. Museum, Jones 75835.
Tribulus ne L:: Calicop. Goat head,
Puncture weed, Abrojo de for amarilla,

Cadillo. NSA; Rabalais (1975)

ACKNOWLEDGMENTS
The authors thank Paul Eubank and Darrell Echols at PINS, M. Bohm
for assistance with the resumen, R. Nelson and M. Goetze for field assis-
tance, as well as Biology Research Problems, Ecology, Field Biology, and
Plant Taxonomy students at TAMUK and TSU for helping with parts of
this investigation. The authors acknowledge support from the Dean’s Shar-
ing Fund at TAMUK and Organized Faculty Research at TSU.

REFERENCES

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SMITH, : ‘ 1996. Ecology and field biology, fifth edition. Harper Collins College Pub-
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1246 Sipa 18(4)

BOOK NOTICE

Roperr W. ALLARD. 1999. Principles of Plant Breeding. Second Edi-
tion. (ISBN 0-47 1-02309-4, hbk.). John Wiley & Sons, Inc., 605 Third
Avenue, New York, NY 10158 (212-850-6336). $69.95. 254 pp.

From the Introduction: “In broadest terms, this second edition of Principles of Plant Breed-
ing is concerned with the two distinct components of Darwinian evolution: (1) the facts of
evolution (“descent with ee and (2) selection as the chief agent of evolutionary
change. However, the narrower focus of the book is on two quite recent and specialized
rluci y s: first, che evolution of crop plants under cultivation, a process that started
about 13,000 to | 1.000 years ago with the cultivation and ultimate domestication of many
wild plant species in various of the world; and second, the current scientific phase of en
Sees. a ae me Seantee cnyt in the twentieth century as Darwinian and M
he framework within which evolutionary ae

lian pr inc ples

in livine paoainae occur.”

Contents of Principles of Plant Breeding: Preface. Part I. Introductory Topics: Darwin-
ian Evolution, Origins of Agriculture, Evolution During Domestication, ae Systems
of Plants, Overview of Plant Breeding; Part Il. Biological Foundations of Plant Breed-
ing: Heredity and Environment, Genetic Ses enee of Hybridization, foheritenee of
Continuously Varying Characters: Biometrical Gen , Evolution During Cultivation,

Marker-Assisted ree. of Adaptedness in Nature, Marker. Assisted Dissection of Adaptedness
in Cultivation. Part III. Modern Breeding Plans: Reproductive Systems and Breeding
Plans, Breeding Self-Pollinated Plants, Breeding Hybrid Varieties of Outcrossing Plants,
Breeding Clonally Propagated Plants, Breeding Hybrid Varieties of Selfing Plants and Plants
that Are Clonally Propagated in Nature, Plant Breeding for Low-Input Agriculture, Glos-

sary, References, and Index

AN ANNOTATED PRELIMINARY CHECKLIST
OF THE DICOTYLEDONOUS LIANAS
AND VINES FROM THE LAS CRUCES
BIOLOGICAL STATION, COSTA RICA

ALEXANDER KRINGS!

Department of Forestry
North Carolina State University
Raleigh, NC 27695, U.S.A.

ABSTRACT
In order to contribute to our understanding of lianas and vines, as well as to facilitate
future research, a preliminary checklist of the dicotyledonous lianas and vines from the Las
Cruces Biological Station, Costa Rica is presented. Seventy species in sixty genera and thirty-
two families are recorded. The largest climbing families at Las Cruces are Cucurbitaceae
(11 spp.), Leguminosae (6 spp.), Sapindaceae (5 spp.), and Bignoniaceae (4 spp.).

RESUMEN
Para contribuir al conocimiento de lianas y enredaderas, asf como para facilitar otras
investigaciones, se presenta una lista preliminar de las lianas y enredaderas dicotiled6neas
de la Estacién Bioldgica Las Cruces, Costa Rica. Se citan 70 especies de 60 géneros y 32
familias. Las familias mas grandes de Las Cruces son Cucurbitaceae (11 spp.), Leguminosae
(6 spp.), Sapindaceae (5 spp.) y Bignoniaceae (4 spp.).

INTRODUCTION

Lianas are important constituents of tropical forests that have, until re-
cently, been largely neglected in both botanical and ecological studies (Gentry
1991). Arguably the most important physiognomic character differentiat-
ing tropical and temperate forests (Croat 1978), lianas are woody vines, beginning
life as terrestrial seedlings and capable of growth in mature forests (Gentry
1991). Herbaceous vines also start life as terrestrial seedlings, but are typi-
cally found in disturbed habitats and lack significant secondary growth. Ninety
percent of the liana species of the world occur in the tropics (Walter 1985).
Vegetation studies of Central and South American forests indicate that li-
anas can occur on 42 to 50 percent of forest trees (Montgomery & Sunquist
1978; Putz 1982, 1984). In the last two decades, various lists covering dif-
ferent aspects of the Costa Rican flora have been published (Janzen & Liesner
1980; Hartshorn & Poveda 1983; Haber 1991; Kappelle et al. 1991), how-

'Present address: Zilker Botanical Garden, 2220 Barton Springs Rd., Austin, TX 78746,
U.S.A.

Sipa 18(4): 1247-1258. 1999

1248 Stipa 18(4)

ever, no list has yet been published treating specifically the liana or vine
flora of Costa Rica. In order to contribute to our understanding of lianas
and vines, and to facilitate future research, the present study sought to document
the dicotyledonous lianas and vines of the Las Cruces Biological Station,
Costa Rica

a site not previously subjected to systematic liana collections.

METHODS
Site Description
The forest of the Las Cruces Biological Station is classified as tropical premontane
rain forest following the Holdridge Life Zone System (Holdridge 1947; Hartshorn
1983). It is a relatively tall forest (30-35 m) with abundant oaks and an
epiphyte load conspicuously less than at similarly classified zones at Monteverde

(Hartshorn 1983; Krings, pers. obs.). The vegetation of the site is still in-
completely known, especially when compared to the more intensively studied
La Selva Biological Station. Hartshorn & Poveda (1983) present a prelimi-
nary list of sixty-three tree species known from the site.

he climate of the Station is illustrated in Figure | by a Walter climate
diagram based on data from the Station’s meteorological station for the years
1996 and 1997. The mean annual rainfall at the Station is 4236 mm and
the mean annual temperature 20.6° C. Stiles et al. (1989) show weather data
from the Instituto Meteorologico de Costa Rica indicating mean annual
precipitation and mean annual temperature for the nearby town of San Vito
to be 3988 mm and 21.7° C respectively.

As seen in Figure |, rainfall is somewhat seasonal with two distinct peaks
from May—June and October-November broken by a ‘verani//o’ from July—
August. December is the only month in which mean monthly rainfall may
drop below 100 mm.

The temperature at the Station fluctuates relatively little throughout the
year. As indicated in Figure 1, the highest mean monthly temperature is
21.8° C and the lowest mean monthly temperature 18.7° C. The highest
recorded temperature for the two years is 32.4° C and the lowest 15.2° C.

COLLECTION AND DEPOSIT

The dicotyledonous lianas and vines of the Las Cruces Biological Station
were collected from August to October 1996 and in March 1997. The col-
lections were made between 1000 m and 1200 m elevation.

Specimens were collected using expandable clipper poles and, in some
cases, by climbing the host tree. Sometimes mountaineering ropes were used
to assist climbing by attaching them to a weighted fishing line and then
shooting the line over a host tree branch with a slingshot (see Perry 1978;
Moffett 1993; Laman 1995). The fishing line was used to pull up parachute
chord, which unlike fishing line, will support the weight of mountaineer-

Krinas, Lianas and vines from the Las Cruces Biological Station 1249

[ LAS CRUCES (1193 m) 20.6° 4236

°C —L
=. 22]
t 500
+ 400
+ 300
. 200

100
32.440
21.89
37 20a . . ; . .
| Sr aa ae aan ae”
Jan Feb Mar Apr May — Jun Jul Aug Sep Oct Nov Dec

Fic. 1. Climate of the Las Cruces Biological Station, Costa Rica as a Walter climate diagram
oe on data from the Station’s meteorological station from 1996-1997).

ing rope. Climbing devices, known as ascenders, were then used to clink

into the canopy where samples were collected. Voucher specimens were dep
at CR and F

RESULTS

Seventy species in sixty genera and thirty-two families are recorded. Taxa
are arranged alphabetically by family, genus, and species. An asterisk pre-
ceding a name indicates that species to be collected from cultivation. An-
notations include the relative rarity (Table 1), habit (liana or vine), diameter
at breast height (dbh) or height climbed when known, and any distinguishing
characteristics. As the relative rarity descriptions are based on casual obser-
vation they are not definitive. A complete census of the lianas and vines at

TABLE |. Relative rarity categories.
Category Definition
Uncommon One to five individuals observed

Common Six or more individuals observed

1250

Stpa 184

—

Las Cruces will most likely change many of these observations. Unless in-
dicated otherwise, notes on distinguishing characteristics come from the
author's own study of field and herbarium specimens. For the purposes of
the annotations, elevations above 700 m are considered ‘montane.’

ACANTHACEAE
Mendoncia brenesii Stand. (Krings 177)—

Uncommon at Las Cruces. Collection taken

from liana climbing to 15 m, 50—60 m from
forest edge at an ae of LL00—1200 m
Thi

US Opposite. -leaved liana i is recognized by th
41
es

subt grooved stems

and Gai ovate bracts. Te fait 3 is bilateral.

Mendoncia tonduzii Turrill (Krimes 213)—
Uncommon at Las Cruces. Collection taken
from liana climbing to 11 mat an elevat

of 1180-1190 m. Recognized “ the 4- ie
hirsute stems and the oblong-elliptic, mu-
cronate bracts, with a rounded apex. The some-

what ae purple fruit is asymmetrica
near the apex

AMARANTHACEAE
Chamissoa altissima (Jacq.) Kunth (Krings
156)—Uncommon at Las Cruces; only one

individual seen. Co
bust liana

lec tion taken from ro-
, 5.7 cm dbh, climbing to 17 m
at an elevation of 1120-1200 m. Relatively

non-descript, alternate-leaved climber with

small flowers subtended by 1—3 bracts in
pyramidal terminal inflorescences.

APOCYNACEAE

AA ] | 1

K. Schum. (Krings
189)—Uncommon at Las Cruces. Collection
taken from vine climbing to only 2 mat an
O-1L095 m. Distinguished
from the other collected Apocynac climber,

elevation of 105

ve cordate leaf bases
and glands on the midrib base above.
Prestonia tsthmica Woodson (Krings 134
Uncommon at Las Cruces; only one individual
seen. Collection taken at 1015 m elevation,
from flexible vine at forest edge with

Prestonia isthmica, by tl

corky,
sticky
exudate, Young leaves velutinous above and
below (see above for other distinguishing

cream-colored older stem and clear,

characters).

BIGNONIACEAE

Amphilophium pannosum (DC.) Bureau
& K.Schum. (Krings 162)—Common at Las
Cruces. Collection taken from liana 4.3 cm
dbh, climbing to 25 mat an elevation of 1075
m. Vegetatively easily recognized by the villous,
sharply hexagonal stems with long trichomes
(to 2 mm), and trifid tendrils. The compressed,
ellipsoid, rugose-tuberculate capsule is also
unmistakable. Separated from Pithecoctenium,

the only other wild Bignon genus in Cost
Rica wit

1 hexagonal stems, by the nde
trichomes of vegetative parts (simple in
Pitheco sia

aea verrucosa on andl.) A.H. Gentry
(Krings 147)—Comm
lection taken from

nat Las Cruces. Col-
wa to 4.4 cm dbh. Dis-
tinguished from A. patellifera and A. florida,
the only two other Arrabidaea species occurring
above 700 m
presence of interpetiolar gland fields, a bi-

elevation in Costa Rica, by the

labiate sige cies greater the yan 1.5 crn and
]

he siti! oe fried ae at Las
Cruces, ee aa pannosum, by the
r fruit, to 2.5 cm wide (ellipsoid in r

eee to 8 cm wide).
Lundia puberula Pittier (Krings 258)—-
Uncommon at Las Cruces. Collection taken
from vine growing over low shrubs at 1075
m elevation. Similar to Arrabidaea, but dis-
tinguished by the pubescent anthers (glabrous
in Arrabidaea) and pubescent to velutinous
capsules (glabrous, or with scattered trichomes,
to tuberculate in Arrabidaea).
Martinella ol | h) Bur. & K. Schum
sks xy 1S8O)—Uncommon at Las Cruces; only

individual seen. Collection taken from
15 m, 50
forest edge at an elevation of 1100-1200 m.

i climbing to )—60 m from the

Gentry (1973) suggests good field charac-
ters include frequently bending or twisting

Krincs, Lianas and vines from the Las Cruces Biological Station

petiolules, trifid tendrils, and conspicuous
interpetiolar ee Unrivaled sl any other
Bignon in Costa Rica, the narrowly linear
fruit (to 1.8 cm oo. can ener 130 cmin
length

BORAGINACEAE

Tournefortia sp. (Krings 142)—Uncommon
at Las Cruces. Collection taken from liana
climbing into canopy at the forest’s edge at
1060 m elevation. Somewhat non-descript,
but recognized by the scorpiod inflorescences
and wintergreen odor to the crushed leaves.

CAMPANULACEAE

Burmeistera cyclostigmata Donn. Sm.
(Krings 179)—Common at Las Cruces. Col-
lection taken from vine climbing to 4 m, 50—
60 m from the forest ne cuse at 1100-1200 m
elevation. Recognized by the milky latex,
purplish corolla, a serrate- nae calyx.
pee se es C. Presl (Krings

72)—Common at Las Cruces. Collection
se from vine ae ng to3 mat 1050 m
elevation. Recognized by the bright rec
rolla with yellow lobes.

COMBRETACEAE

CO=

Combretum laxum Jacq. (Krings 165)—
Uncommon at Las Ces: Collection taken

ro I py in clos
ae at 1200 m elevation. Recognized by
the opposite leaves, parallel, tertiary leaf
venation, and distinctive four-winged fruit.

COMPOSITAE

Mikania guaco Bonpl. (Krings 186)—Un-
common at Las Cruces. Collection taken from
liana climbing to 10 m in closed f
1050-1095 m elevation. Recognized by the
opposite leaves, three-veined from the base,
and the glabrate, fistulose stems

Mikania skutchiiS.F. Blake (Krings 166)—
Uncommon at Las Cruces. Collecti

orest at

on taken
from liana climbing into canopy of closed
forest at 1200 m Distinguished
from M. gwaco by the densely villous leaves
and the villous, never fistulose stems

Otopappus verbesinoides Benth. (Krings
197)—Common at Las Cruces. Collection

elevation.

1251
taken from liana climbing to only 1.5 m at
1075 m elevation. Characterized by asperous
stems and opposite leaves (narrower than in
the Mikania spp. above or
below),

the Siuclairia sp.
three-veined from the base. See
Hartman & Stuessy (1983) for a revision of
the gen
Sinclii sass lati (Klatt) Rydb. (Krings
mon at Las Cruces. Collection
wie ae liana climbing to 7 mat 1050—
1095 m elevation. Recognized by the ser-
rate, opposite leaves, three-veined from the
base, light green above, and glaucous below.

CONNARACEAE

Rourea sp. (Krinmgs 203)—Uncommon at Las
Cruces. Collection taken from liana climb-
ing to 7 mat 1170 m elevation. Recognized
by cylindrical pulvini, pinnately compound
leaves, with the basal leaflets alternate to sub-
opposite, and reflexed branchlets aiding in
climbing. Rowrea has 2 carpels (1 in Connarus)
and imbricate sepals (valvate in Cnestidium)
Woodson et al. 1950).

—

CONVOLVULACEAE

Maripa sp. (Krings 277)—Uncommon at Las
Cruces. Collection taken from vine climb-

ing to 17 m at the edge of a forest gap at
1030 m loa Recognized by the rounded
leaf-bases (cordate in Merremia), pnaees
late, whitish-purple flowers, and often im-

ce Merremia 2 fruit

pressed venation. Unli
is indehiscent
Merremia sp. (Krings 21 1)—Uncommon at
Las Cruces; on seen in one location. Col-
lection taken fr

t 1200 pa ies Recognized by the dis-
tinctive large, rounded-squarish seeds and
deeply lobed leaves with
Convolvulaceous venation. Unlike Maripa,
the fruit is dehiscent

m seedlings on forested ridge

typical

CUCURBITACEAE

*Cionoscyos macranthus (Pittier) C. Jef-
as es 280)—Onl
vat t Las

known from culti-
Cruces. Cells tion en aoe

vine NCal the Stati OL

t 1095 aeacen Recognized “ the

1252

pate nioEm glands acai near the leat base

|
below flexuous.

The genus is closely clued to ae oe and
essentially only differentiated by the fleshy,
peponiform fruit (baccate and fibrous in
Cayaponta)
*Cucurbita moschata (Duchesne ex Lam.)
Duchesne ex Poir. (Krings 279)—Only known
from cultivation at Las Cruces. Collection taken
from vine growing in Station vegetable ies
at 1095 m elevation. Recognized by 4—
S-branched tendrils and ee. irritat-
ing leaf hairs.
Cyclanthera multifoliolata Cogn. (Krings
269)—Common at Las Cr
taken from vigorous vine taking over a large

uces. Collection

tree- fall Bap at eae m ae Easily

wlnis ring. U wigue among Costa Rican Cy ees
s patelli-
OED & — near Ge leat base ee

li en.) Pittier (Krines
175)—Uncommon at Las Cr uces. Collection

species in the pre numer

taken from vine growing outside the Station,
on roadside at LOO0—1100 m. Easily recog-
nized by the glabrous to puberulous stems,

3-lobate to angulate leaves, whitish-green
campanulate flowers with fused filaments and

vertically plicate anthers, and smooth, non-
echinate, explosively dehiscent fruit.
Fevillea cordifolia L. (Krings 272)—Uncom-
mon at Las Cruces; only one individual seen.
Collection ce n from young liana —
on the edge of a ridge-top gap at 1200
Rican Cu.

elevation. One of the few Costa

curbits becoming a nop y haps —
| VeSo tatively by tl

curling aleve and below the bifurcation. Dif-

ferentiated from Sicydivm, the only other
montane (1.e., > 700 m) Costa Rican Cucurbits
exhibiting tendrils curling above and below
the bifurcation, by the woody habit, five sta-
mens, and 3-locular ovary (herbaceous, three
stamens, and unti-locular in Sécydiam).

makoyana (Lem.) Cogn. (Krings
276)—Common at Las Cruces. Collection

Gurania

ot

taken from robust vine climbing to 17 ma
the forest edge bercween 1000 and 1100 m
elevation. Gurania is one of only two o gen-
era recognized by simple tendrils, inflorescences

Sipa 18(4)

consisting of several flowers, and orange to
reddish corollas, but can be distinguished by
the orange or reddish calyx lobes (green in
Psigurid). Bepaetes from other Costa Rican

Guranta species by the simple leaves, pedicels

of staminate flowers | to 3 mm long, stem
hairs (if presence) tess than 4 mm long, and
the calyx lobes frequently 2 to 3.5 times as
long as the calyx tube

Melothria sp. (Krings 206)—Common at Las
Cruces. Collection taken from vine climb-
ing to 2 mat 1145 m elevation. Three spe-
cies of Me/othria are known from Costa Rica.
| are quite similar and cannot be conclu-

nd

72)

vely separated vegetatively. M. du/cis is
distinguished by a white corolla and yellow
to orange fruit. Both M. scabra and M. pendula
exhibit a yellow corolla and are differenti-
ated by the mature fruit (at least 2.5 cm long,

light and dark green ane in au nae versus

=
=

ong, gree dla)
Psiguria triphyll a (Miq,) pie (Krings
246)—Uncommon at Las Cruces. Collection

only to 2cm

taken from vigorous vine climbing to 15 m
at LOOO—1 100 related
to Gurania (see G. makoyana for discussion).
Separated from P. warscewiczii, the only other

1 elevation. Closely

Psiguria species above 700 m in Costa Rica,
by the calyx not green-spotted and the co-
riaceous leaves (green-spotted calyx and
membranaceous leaves in P. warscewiczit).
Rytidostylis carthaginensis (Jacq.) Hook.
& Arn. (Krings 250)—Uncommon at Las
Cruces. Collection taken from slender vine
growing on a streamside shrub at 1000-1 100
m elevation. The only montane Costa Rican
Cucurbit displaying an elongate, narrowly
cylindrical calyx tube (to 3 cm long
*Sechium edule (Jacq.) Sw. (Krings 278)—
Known only from ee at Las Cruces.
Collection taken from vine growing in the
1095 m eleva-
ne genus is recognized by tendrils 3—
nectaries at the base of
by the
nectaries sunken into the
the hypanthium (but not protrud-
ing conspicuously below) and the filaments

Station vegetable garden at

—

tion. T
S-fid an
the hypanthium. S. edw/e is recognizec
combination of the

nd ten floral

ion

base of

only partially connate, the anthers free.
Selysia prunifera (Poepp. & Endl.) Cogn.

KrinGs, Lianas and vines from the Las Cruces Biological Station

(Krings 155)—Uncommon at Las Cruces. Col-
lection taken from stout liana, 3.2 cm in
diameter growing into canopy at 1120-1200
m elevation. The only species of Se/ysta in
Costa Rica, it is distinguished from mem-
bers of Cayaponia, Cionosicys, Cyclanthera, and
Tecunumania, the only four other Costa Rican
ibiting patelliform glands on the
abaxial afeieace by the glands frequently
scattered throughout the abaxial leaf surface
(not ee crowded near the pase), the anthers

]
rhe OL merely slightly CULVCd

essentially s Stl al

at the ends, and the arrow-head shaped seeds

DICHAPETALACEAE

Dichapetalum nevermannianum Stand.
(Krings 151)—Uncommon at Las Cruces.
Collection taken from liana to 2.5 cm dbh,
climbing to 17 m at 1075 m elevation. Dis-
tinguished from other Mesoamerican Dicha-
petalum species by the linear-oblong to oblan-
ceolate leaves to only 3. cm wide and the hispid
stems and inflorescences (Lundell 1966).

ERICACEAE

Psamissia ramiflora Kl. (Krings 160)—
Uncommon at Las Cruces. Collection taken
from shrubby liana climbing to 17 mat 1120—

200 m elevation. The only lianescent rep-
resentative of Ericaceae observed at Las Cruces,
P. ramiflora can be recognized by the alter-
nate, plinerved leaves, with swollen petioles.

EUPHORBIACEAE
Dalechampia cissifolia Poeppig wales
mii ena at La s Cruces. Collect
taken from vine ¢ x over low ae
at 1030 m joan rela by the ser-
rulate, trifoliolate leaves, much resembling
Cissus, but se tendrils. Only two other
Dalechampia species in Costa Rica are also
trifoliolate. D. pie has 6 female sepals
and involucral bracts 1O—14 mm long and,
though also displaying 7-11 female sepals
and involucral bract less than 5 mm long,
.) lypt, rey Are | ‘ ] ] bat ps [ee | ]

intermixed on the stems (Burger & Huft 1995).
Manihot brachyloba Muell. Arg. (Krings
7173)—Uncommon at Las Cruces. Collection

1253

taken from scrambling liana climbing to 6
m trailside at 1060 m elevation. Distinguished
from other Costa Rican Manihot species by
the climbing habit and the tri-lobed leaves
(Burger & Huft 1995).

GESNERIACEAE

Capanea sp. (Krings 214)—Uncommon at
Las Cruces? Collection taken from lianescent
plane climbing to 8 mat 1100-1125 m eleva-

nA neisetctices by asynmneenc, apposite

HIPPOCRATEACEAE

Salacia petenensis Lundell (Krings 182)—
Uncommon at Las Cruces. Collection taken
from robust liana to 7.2 cm dbh, climbing
to 20 mat 1060 m elevation. The only liana
observed at Las Cruces climbing by stout,
leafless, curving branchlets (to 1.5 cm diam.).
The | } i | fagiec r le ore eee

HY DRANGEACEAE

Hydrangea peruviana Moric. (Krings 167)—
Common at Las Cruces. Collection taken from
liana climbing with adventitious roots into
canopy at 1190 m beevaion: PecOe nized by

the opposite

brownish

stems with erie ridges.

LEGUMINOSAE

Caesalpinia urophylla (Donn. Sm.) Stand.
(Krings 164)—Common at Las Cruces. Col-
lection taken from liana to 3.3 cm dbh, climb-
ing to 15 mat re m eevalioh: C. wraphy Ha
} | ] ]
is heavily armed
along the stems and leaves. Also distinguished
from other leguminous climbers at Las Cruces
2 the cael bi- planer y compound leaves
and the sp See

pt ic kles both

in more eee, 1 areas in the forest.
Canavalia oxyphylla Stand|. & L.O. Williams
(Krings 191)—Common at Las Cruces. Col-
lection taken from vine climbing to 10 m
at 1015 m elevation. This crifoliolate vine
with pinkish flowers is recognized in fruit
by an additional suture displaced from the
ones on each valve, often appearing medial
on the valve.

1254

Machaerium cobanense Donn. Sm. (Krings
34)—Common at Las Cruces. Collection
fe from liana growing into canopy along
trailside at 1015 m elevation. Recognized
by the paired spines of stems and the rounded
oblong to elliptic leaflets lacking acuminate
tips. Stems may exude a red sap when cut
Machaerium seemannii Benth. ex Seem.
(Krings 193)—Uncommon at Las Cruces. Col-
lection taken from liana climbing to 6 m at
1030 m elevation. Recognized by the lan-
ceolate leaflets with acuminate tips and the
leading shoots often leafless, but armed with

paired spines, and curling much like tendrils.
Mucuna sp. (Krings 135)—Common at Las

uces. Collection taken from vine in forest
interior at LO15 m elevation. Generally, unmis-
takable lianas due to the long pendent flowers
and fruits, both often with urticaceous hairs.
Rhynchosia erythrinoides
(Krings 149)—Common at Las Cruces. Col-
lection caken from liana with flattened stem
gto 15 mat 1075 melevation. Rec-
cnet vegetatively by the somewhat rhombic

Cham. & Schltdl

limbin

leaflecs of the trifoliolate leaf and the flattened

stem (no other liana at Las Cruces has been

observed with such a stem).
LOGANIACEAE

Strychnos sp. (Krings 183)—Common at Las
Cruces. Collection taken from liana climb-
ing to 6 mat ued m veevadion: Easil y dis-

tinguished fr leaved,
tendrillate ae Hes in co Rica,
Bignoniaceae, by the simple, often 3-veined

leaves. Tendrils in Strychnos also tend to be
more hook-like

MALPIGHIACEAE

Hiraea hte Standl. & L.O. Williams
(Krings 148)—
Collection taken from liana climbing to 20
mat 1075 m elevation. Recognized by densely

ommon at Las Cruces

ferruginously tomentose stems and samaras
with semi-circular lateral wings and reduced
dorsal wings. Stipules are often borne above
the petiole base

Tetrapterys sp. (Krings 170)—Uncommon

at Las Cruces. Collection taken from liana

Stipa 18(4)

climbing to 25 mat 1060 m elevation. Rec-
ognized by samaras with four oblong, elon-
gate lateral wings and reduced dorsal wings.

MARCGRAVIACEAE

Marcgravia sp. (Krings 273)—Uncommon
at Las Cruces. Collection taken from mate-
rial off the forest floor on a ridge at 1200 rm
elevation as the high-climbing liana was un-
reachable. A very distinct adventitious root
climber, recognized when young by the of-
ten angular stems growing appressed to the
trunk of trees wah the leaves te flac.

The alternate, fre quently

dark-punctate leaves, as well as the whorl of
flowers harboring a whorl of saccate necta-
ries in its center, are also unmistakable.

MENISPERMACEAE

Anomospermum reticulatum (Mart.)
Eichler (Krings 184)—Uncommon at Las
Cruces. Collection taken pom robust liana to

5.3m dbh, chnaPing to4 mat 1030 m eleva-

] 13
unifolio-

tion. S
late cece ate to the sv enen pulvini, A.

] }
1 ecognized WLICKI

reticulatum
dried by the glossy, reticulate leaf venation.
Cissampelos pareira L. ngs As eas

mon at Las Cruces. Coll vine
climbing to 4 mat 1030 m sevavion: The
genus Cissampelos in our area is distinguished
from other tropical Menisperm genera, such
as Anomospermum, Chondrodendron, H id

Odontocarya, by the staminate flowers with
n the afore eres

anc

= pL

nly 4 sepals
ae and the nedtacents Talste (Ex xdes
C. parerra is distinguished from C. nel
in that the leaves are not peltat
Cissampelos tropaeolifolia DC. (Krings
268)—Uncommon at Las Cruces. Collection
taken from vine growing over low shrubs on
trailside at 1150 m elevation. A mor
weedy vine, C. tropaeolifolia is distinguished
from C. pareira by the peltate leaves and seri-

or less

ceous indument of the leaves.

NYCTAGINACEAE

Pisonia aculeata L. (Krings 168)—Uncommon
at Las Cruces. Collection taken from stout

KRINGs, Lianas and vines from the Las Cruces Biological Station

liana to 5.2 cm dbh, climbing to 20 m at
1190 m elevation. This opposite-leaved li-
ana is armed with stout, axillary, recurved
spines and is recognized in fruit by the stalked

glands arranged linearly along the angles of

the fruit.

PASSIFLORACEAE

Passiflora costaricensis Killip (Krines 27 1)—

Las Cruces. Collection taken
1: | EG

Uncommon at
from a small liana
thickets at 1170 m elevation. Distinguished
from the other two collected Passiflora species,
| } | Ess [eee ] ] } ] B ]
my

g through trailside

] : ] Disti ey | 1f }

bilobate montane hace species, by the
leaves not peltate, 1 1/2 times as long as wide
(or nearly so), and dine ae leaf lobes gen-
erally 1/3 the length of the midrib (1/2 or
more in P. capsularis). The fruit is reddish,
ellipsoid, and asymmetrical in cross-section.
Passiflora menispermifolia Kunth (Krings
150}—Uncommon at Las Cruces. Collection
taken from a small vine climbing to 4 m in
the understory at 1075 m elevation. Distin-
guished from other Costa Rican Passiflora
species with trilobate leaves by the leaves
essentially entire, not peltate, not glandu-
lar-ocellate below, villous with straight hairs
(puberulent with hooked hairs in P. /obata),
and petioles with 2 or more gland pairs.

Passiflora oerstedis Mast. (Krings 215)—
Common at Las Cruces. Collection taken from
vine climbing to 4 m in understory between
1100-1125 m elevation. Distinguished from
other montane Costa Rican Passiflora species
with unlobed leaves and cordate leaf bases,
by the petioles ee 3¢ or more, linear to alot,

glan d pairs,

(parallels in P. le

PIPERACEAE

Sarcorhachis naranjoana (C. DC.) Trel.
(Krings 144)—Common at Las Cruces. Col-
lection taken from liana climbing to 6 mat
au m elevation: Recognized by alternate,
swollen nodes, and solitary.

axillary, spicate inflorescences (leaf-opposed

——

in Piper)

1255

RANUNCULACEAE

Clematis dioica L. (Krings 195)—Uncom-
mon at Las Cruces. Collection taken from vine
climbing to 5 m at 0 m elevation. Eas-
ily recognized by the opposite, pinnately-
compound leaves (3—5 leaflets) and the ra-
g around objects, much like
a tendril, while lisbine.

ts often curlin

RHAMNACEAE

Gouania sp. (Krings 138)—Common at Las
Cruces. Collection taken from sterile vine
growing in forest edge tangle at 1015 m
elevation. Recognized to genus by the ten-
dril axillary toa terminal leaf or inflorescence
at apex of short branch and often coiled like
a butterfly proboscis. Leaves usually have three
basal veins. The collection is probably either

(ee or G. Plies the former dis-
tinguished by oral disc somewhat pu-
ees nt with eet long, conspicuous
trichomes (glabrous or minutely, appressed
puberulent in G. /wpuloides).

RUBIACEAE

Manettia sp. (Krings 264)—Common at Las
ruces. Collection taken from young vine
growing over low shrubs in a forest gap at
1070 m elevation. These herbaceous vines
are recognized by the opposite leaves with
distinct arcuate venation. Corollas are tubular
to salverform
ndia err Lorence & Dwyer (Krings
90)—Uncommon at Las Cruces. Collection
taken from liana climbing to 12 mat 1050—
1095 m elevation. Recognized by the sub-
terminal, more or less stout, spines in groups
of 3 to 4. Uncaria spines occur in pairs at
the nodes.

SAPINDACEAE

eee alata (Ruiz & Pav.) G. Don (Krings

0)—Common at Las Cruces. Collection
ee from liana growing into canopy at 1030
m elevation. Distinguished from other pin-
nately compound-leaved montane Pau/linia
species by the leaves 2-jugate, the leaflets of
the lowest pair simple stipules only to 3mm

1256

long, the unwinged, red fruit to 2 cm long,
and the stem cross-section showing 3—6
peripheral vascular cylinders surrounding a
central one,

Paullinia bracteosa Radlk. (Krings 145)—
Common at Las Cruces. Collection taken from
stout liana to 4 cm dbh at 1075 m eleva-
tion. Easily distinguished from other pin-
nately compound-leaved montane Pau/linia
long)
and the stem cross-section showing a single

species by the large stipules (2-5 cm

vascular cylinder.
?aullinia grandifolia Benth. ex Rac
163)—Common at Las Crv
taken trom liana climbing into canopy at 1180
m elevation. Distinguished from P. mallophylla

Qu

Ik. (Krimes

ices. Collection

P. ingaefolia, the only other montane

Paullinia species with the lowest leaflets of

3-5 jugate leaves trifoliolate or pinnate,
by tl I fruit Qwinged in P. mellophylla)

by
caieeuily triangular stipules, to 8 mm
g (ovate to broadly lanceolate, 1.5 to 4

cm = eae in P. ingaefolia

Paullinia pterocarpa Tana Pinch (Krings

8)—Uncommon at Las Cruces. Collection
taken from liana to 2.6 cm dbh, climbing
into canopy at 1120-1200 m elevation.
Distinguished from other winged-fruited
montane Paullinia species, by the pinnately
compound leaves with the lowest leaflet pair
simple (all others are either trifoliolate or with
the lowest Le oe sees te).
nes 154)—Uncom-

mon at Las Cruces. C ac from liana to

Serjar La

3.2 cm dbh, climbing to 15 m into canopy at
i 1 20-1200 m elevation. Distinguished from
S. lobulata, the only other pinnately 5-foliolate
montane Serjania species, by the densely hirsute
stems and petioles (hairs 1-2 mm long) and
che conspicuous stipules, to 5 mm long (in-

2

conspicuous, to 3. mm in S. /obulata).
SOLANACEAE

Solanum sp. (Krigs 176)—Uncommon at
Las Cruces? Collection taken from lana climb-

SIDA 18(4

Ww

ing near forest edge at 1 100-1200 m eleva-
tion. The genus is recognized by calyces with
5 vascular ribs or lobes and anthers dehisc-

ing by terminal pores (D’Arcy 1973).
ULMACEAE

Celt tis igu andae@ea (Jac q VS

Sarg. (Krings 181)—
Wacommen at Las Cruces. Collection taken
from robust liana to 4.3 cm dbh, climbing
to 10 mat 1025-1075 m elevation. Although
; : ad oe

Li

(se errulate to serrate at tip, 3-veined from base),

a

. iguanaea is easily recognized by the lack
of tendrils, the presence of spines, the more

r less asymmetrical leaf bases, and the asperous
leaf surfaces.

VITACEAE

Cissus rhombifolia Vahl (Krinzgs 192)—
Commonat Las Cruces. Collection taken from

vine climbing to 3 mat 1060 m elevation.
Distinguished from other trifoliolate Costa

Rican Cissus species by the more or less rhombic

terminal featee and the small fruits (ca. |
cm long and wide).

Cissus verticillata (L.) Nicolson & C.E. Jarvis
(Krines 141
ection taken from liana growing into canopy

)—Common at Las Cruces. Col-

—_—

at 10 me ‘levation. Perhaps the most common

of the Costa in simple-leaved Cissus species,

C. eee ney is ae qed by the leaves
no tly dimorphic (dimorphic in C,
hiformifolia, broadly ovate (narrowly ellip-
tic to lanceolate in C. as anes the pedicels
glabrous (hircellous in C. cacuminis
hae tiliifolia Humb. & Bonpl, ex Roan,
Schult. (Krings 210)—Common at Las
Ps ruces. Collection taken fi
ana to 7 cm dbh, climbing to 17 mat 1120

rom vigorous li-

m elevation. The only representative of Vitis
in Costa VY. teliifolia is distinguished
from Cissus by the 5-merous flowers, the panicle

inflorescence, and the leaves densely floccose-
tomentose beneath.

DISCUSSION

This list should be treated as a working checklist. Included are only col-
lections that have been assigned with confidence to either genus or species.

KRINGs, Lianas and vines from the Las Cruces Biological Station ay

A complete collection list, including unknowns, has been deposited at F.

Nine of the eleven tendrillate climbing families of Costa Rica (see Krings
1997), are represented at Las Cruces, although the Leguminosae are repre-
sented by only non-tendrillate species. No climbing, tendrillate Polemoniaceae
(Cobaea) or Polygonaceae (Antigonon, introduced) have been found. This is
not surprising as Cobaea vines are generally found above 1600 m in Costa
Rica, well beyond the upper limits of Las Cruces. Only C. gracilis and C.
scandens are known to have been collected as low as 700-1000 m (Krings
1997)

Although incomplete, I hazard that the list includes the majority of the
lianescent taxa of Las Cruces. Currently, the largest climbing families are
Cucurbitaceae (11 spp., incl. 3 cultivars), Leguminosae (6 spp.), Sapindaceae
(5 spp.), and Bignoniaceae (4 spp.)—together comprising 37.1 % of the
recorded species. Most additional species records are likely to come from
the following, mostly vining families: Asclepiadaceae, Convolvulaceae,
Loganiaceae, and Solanaceae. Among monocotyledons, more work is needed
for all taxa, but especially Smilax L. and Dioscorea L., both of which have
been seen on the grounds.

ACKNOWLEDGMENTS

This study was made possible through a travel grant and assistantship
provided by the College of Forest Resources at North Carolina State Uni-
versity, fellowships in Tropical Biology from the Organization for Tropical
Studies (OTS), and a visiting scientist scholarship from the Field Museum
of Natural History. I thank OTS for permission to conduct research at Las
Cruces and for help with collecting permits. In addition, I thank my advi-
sor, Richard Braham, for his insight, encouragement, and support of the
project. I also thank Luis Diego Gémez for his support and insightful com-
ments. As well, I appreciate very much the help and support of the follow-
ing folks: William Burger; the curators and staffs of CR, DUKE, F, INB,
MO, and USJ; James Hardin; Gail Hewson de G6mez; Richard Lancia; Silvia
Lobo; Ronald Murillo; the friendly staff of Las Cruces; and Wendy Worley.
I thank William Burger and an anonymous reviewer for their thoughtful
comments on the manuscript.

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GEntRY, A.H. 1973. Flora of Panama: Bignoniaceae. Ann. Missouri Bot. Gard. 60:781—977.
Gentry, A.H. 1991. Distribution and evolution of climbing plants. In: KE. Putz and

1258 Stipa 18(4)
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34:63-120.
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10:1

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55-157.

GAMOCHAETA SIMPLICICAULIS (ASTERACEAE:
GNAPHALIEAE) IN FOUR SOUTHEASTERN
STATES AND NEW FOR NORTH AMERICA

GUY L. NESOM
Biota of North America Program—North Carolina Botanical Garden

University of North Carolina
Chapel Hill, NC 27599-3280, U.S.A.

ABSTRACT

The presence of the South American native Gamochaeta simplicicaulis (Willd. ex Spreng.)
Cabrera is documented for North Carolina, South Carolina, Florida, and Alabama, based
on specimens in NCU and US. It has not previously been included in accounts of the North
American flora. The species apparently is abundant on the coastal plain of the Carolinas,
where first collected in 1957, and may be expected to occur with frequency in other re-
gions of the Atlantic and Gulf Coastal plain. A morphological description, illustration,
and specimen citations are provided.

RESUMEN

Se documenta la presencia de la planta nativa de Sur América Gamochaeta se ae ee
(Willd. ex Spreng.) Cabrera en Carolina del Norte, Carolina del Sur, Flor labama
basada en especimenes depositados en NCU y US. No habfa sido ae a ipiey eadente en
los informes de la flora norteamericana. La especie es aparentemente abundante en la Ilanura
costera de las Carolinas, donde fue colectada por primera vez en 1957, y puede que se dé
con frecuencia en otras regiones de la Ilanura costera del Atlantico y del Golfo. Se ofrecen
una descripcién morfoldgica, una ilustracién y citas especimenes

The South American native Gamochaeta simplicicaulis (Willd. ex Spreng.)
Cabrera was noted to occur in Florida by Pruski (1997) in his account of
the Asteraceae of the Venezuelan Guayana. The distribution of this species,
otherwise unreported for North America, is further documented here for
North Carolina, South Carolina, and Alabama, based on specimens housed
in NCU. It was not included in accounts of Gamochaeta (Nesom 1990) for
the United States or the southeastern United States (Godfrey 1958; Arriagada
1997). Gamochaeta simplicicaulis apparently is relatively widespread on the
coastal plain of the Carolinas, where it is known from five coastal and near-
coastal counties of North Carolina and every South Carolina county adja-
cent to the Atlantic Ocean, except one. The localities in Walton Co., Florida
(eastern panhandle), and Mobile Co., Alabama, are immediately adjacent
to the Gulf coast.

The first collections of Gamochaeta simplicicaulis from the United States

Sipa 18(4): 1259-1264. 1999

1260 Sipa 18(4)

apparently were made in 1957 from six South Carolina counties in connec-
tion with preparation of the Flora of the Carolinas. Judging from the al-
ready relatively abundant distribution of the species by the mid 1960s, at
least in che Carolinas, it is probable that its current distribution is broader
than the present report indicates. The species almost certainly occurs 1n Georgia
and should be sought in Mississippi and Louisiana. It has been collected
from one county nearly contiguous with southeastern Virginia (Pasquotank
Co., NC).

The native range of Gamochaeta simplicicaulis is South America, where it
is known from Argentina, Chile, Bolivia, Uruguay, Paraguay, Brazil, Guyana,
Venezuela, and Colombia (Cabrera 1961, 1963; Pruski 1997). It also has
become naturalized in New Zealand (Drury 1971; Webb 1988).

Gamuochaeta simplicicaulis in NCU collections has been identified as Gamochaeta
purpurea (L.) Cabrera and Gamochaeta americana (P. Mill.) Wedd. but is con-
sistently distinguished from both, as in the following key.

1. Upper surface of leaves loosely and lightly but persistently tomentose, dull;
involucral bracts lightly villous-tomentose on the lower third; biennial.

G. purpurea
1. Upper surface of s completely glabrous, shiny; involucral bracts glabrous

from base to apex; annual or biennial. 2
2. Plants (30—-)50—-85 cm tall, usually basally and strictly erect; basal and
lower cauline leaves on elongate internodes, commonly withered and de-
ciduous at flowering, relatively even-sized to above midstem, the upper
cauline linear-lanceolate to linear-oblanceolate and apically long-acute,
clusters of small leaves produced in the cauline axils; capicula in small
glomerules in an elongate, interrupted capitulescence, sometimes with

well-developed lateral branches, the puoi rules usually subtended by linear,

spreading bracts longer than the glomerule involu

—

ucral bracts ovate,
apically acuce-acuminate, hyaline aicvout eaibk annual or biennial, flowering
mostly July-August

. simplicicaulis
2. Plants 15—3 550) cm tall, ie decumbent-ascending; basal leaves in
a rosette, persistent at flow , the cauline leaves quickly reduced in
SIZe ee the bas , the ure most ob lanceolate and lh rounded,

axille Ary | 1 1 Narrow, € slong ate genera illy
compact but sometimes interrupted piece. the bracts few and
shorter than the glomerules; outer involucral bracts elliptic-ovate, apically

obtuse to cai often slightly purplish; biennial, flowering mostly April-

June. G. americana

Gamochaeta simplicaulis (Willd. ex Spreng.) Cabrera, Bol. Soc. Argent.
Bot. 9:379. 1961. (Figs. 1, 2) Gnaphalium simplicicaule Willd. ex Spreng., Syst
Veg. 3:481. 1 he linea iat var, pe ee eae Id. ex Spreng. Klatt,
Linnaea 42:140. 18 pe: {Venezuela. Edo. Sucra.}] Cumand, Humboldt 343 (B-
Willdenow fic ‘a 15: oo. ered U an

al

Annuals or biennials, strictly erect, (30—)50—85 cm tall, fibrous-rooted,
the stems commonly simple, strictly erect, and single from the base or with

Nesom, Gamochaeta simplicicaulis

Fic. 1. Habit and involucre of Gamochaeta simplicicaulis (Ahles 32138).

1262 Stipa 18(4)

3mm

Fic. 2. Involucral bracts of Gamochaeta simplicicantis (A hles 30684) inner, middle, and outer

(from left to right).

2—5 basally ascending stems, densely and closely white-tomentose. Leaves
densely and closely white-felty-matted beneath, completely glabrous and
shiny above at maturity, flat, the midrib not depressed, with margins closely
undulate and appearing nearly crenulate, the basal and lower cauline leaves
oblanceolate to oblanceolate-spatulate, 5—9 cm long, 0.6—-1.8 cm wide, on
somewhat elongated internodes (not in a rosette), the basal leaves usually
withered and deciduous by flowering, gradually becoming smaller above
mid-stem, the upper cauline leaves linear-lanceolate to linear-oblanceolate
and apically long-acute; clusters of small leaves produced in the axils of mid
and upper cauline leaves. Capitula in small glomerules in a narrow, elon-
gate, interrupted capitulescence (8—)16—30 cm long, sometimes with well-
developed, ascending, lateral branches, the glomerules usually subtended
by linear bracts longer than the glomerules and usually spreading at right
angles; involucres cylindric-campanulate, 3—3.5 mm high, the involucral
bracts ovate to oblong, scarious and brownish to tan, without red or purple
pigments, glabrous from base to tip, apically acuminate-apiculate. Achenes
0.5—0.6 mm long. Flowering June—)July—August(—October).

Gamochaeta simplicicaulis apparently grows in open sites, as the habitats
are described from the southeastern US, but they otherwise are variable (roadsides,
fields, open woodlands, dunes), except that a sandy substrate may be con-
stant. Pruski (1997) noted that habitats for the species in the Venezuelan

Nesom, Gamochaeta simplicicaulis 1263

Guayana are “savannas, roadsides, forest borders, tepui meadows, rock out-
crops, 800-2100 m.” The late summer to fall flowering of G. stmplicicaults
also distinguishes it from all other eastern North American Gamochaeta, which
flower primarily in the spring.

Collections examined. UNITED STATES. ALABAMA. Mobile Co.: Dauphin Island,
Beauregard St., near Apalache Ave., sunny roadside, 15 July 1965, R. Deramus D576 (NC
FLORIDA. Walton Co.: edge of pine woods along Laird Drive, 4 mi S of sa are 5 Jul
1971, H.A. Davis 15917 (US fide John Pruski). NORTH CAROLINA. Brunswick Co.:

along railroad tracks 1/2 mi S of US routes 74 and 76, 25 Jul 1965, MLN. Sears oid (NCU).

ew er Co.: raised land in brackish marsh, causeway to Wrightsville Beach, 29
Jun 1963, 4. E, Ahles 58631 (NCU); dredge island on Wrightsville Causeway, sand, 29
Jun 1963, A. McCrary 405 (NCU). Pasquotank Co.: vicinity of Weeksville on NC 170,
disturbed swamp border, 25 Jun 1958, H.E. Ables 44345 (NCU). Pender Co.: ca 5 mi N
of jct Hwy 210 and Hwy 17 in Hampstead, on Hwy 17 at St. Jude’s church, Ree field
with scattered pines, eee oO G. Netom NC997-5 (BRIT, FLAS, GA, GH, LSU, MISS,
NCU, TEX, US, USCH — to be distributed). Wayne Co.: Cliffs of the Ness State
Park, open area through eae 10 Jul 1965, H.E. Ahbles 59659 (NCU). SOUT
LINA. Berkeley Co.: 3.5 mi S of jct SC 6 ind 311 on dirt road (W of oak Corner),
thickets, 5 Jul 1957, H.E. Ahbles 30684 (NCU-2 sheets); ca 2 mi W of Honey Hill on SC
45, sandy roadside, 15 Jul 1959, H.E. Ab/es 52983 (NCU); waste ee around the Santee
Wool Combing Mill, Jamestown on SC Rte. 45, 11 Jul 1960, Ables 53783 (NCU).
Charleston Co.: 3.3 mi from Warrens Cross Roads on county roe ai 21 Jul 1957, HLE.
Ables 32138 (NCU). Dorchester Co.: 1.5 mi ESE of Bamberg-Dorchester Co. line on County
11, 20 Jul 1957, H.E. Ables 31825 (NCU). Georgetown Co.: Retreat Beach, Sand Flat,
22 Aug 1957, A.E. Radford 28509 (NCU); Du Bordieu Island, inner sand dunes, 22 Jul
1961, H.E. Ahles 54865 (NCU). Horry Co.: SSE of Conway at jct 4.3 mi ESE of PeeDee
Crossroads, low woods and pinelands, 19 Oct 1957, C.R. Bell 10930 (NCU). Jasper -
Savannah, N.W. Refuge, 28 Aug 1959, E.0. Me/linger s.n. (NCU). Orangeburg Co.:

mi W of Orangeburg- sea county line on US 178 and 2.3 mi SW of dirt road, ee
upland woods, 19 Jul 1957, H.E. Ables 31807 (NCU). Williamsburg Co.: 5.6 mi S of
Heminway on SC 41, sandy near e, 15 Jul 1959, H.E. Ahbles 52984 (NCU).

The NCU collections were variously annotated in 1997 as Gamochaeta
purpurea and G. americana by Dr. J.E. Arriagada, as part of the complete
NCU collection of southeastern US Inuleae (sensu /ato) studied for the Ge-
neric Flora of the Southeastern United States (Arriagada 1997). His anno-
tations for the entire set, however, without exception, merely repeat the
most recent previous identification, except to update the generic name or
replace putative synonyms. A discussion of unresolved taxonomic and no-

menclatural complexities regarding G. purpurea and G. americana, as well
as other previously unreported species of Gamochaeta for the North Ameri-
can flora, is forthcoming (Nesom in prep.

ACKNOWLEDGMENTS

Tam grateful to John Pruski (Dept. of Botany, Smithsonian Institution)
for providing collection data and a photocopy of the Florida collection at
US (first identified correctly by Harold Robinson), as well as various com-

1264 Sipa 18(4)
ments on the manuscript, Ted Barkeley and Lowell Urbatsch for comments
on the manuscript, Jim Massey and Mary Felton (NCU at UNC-CH) for
help in herbarium studies, Bill Burk and Jeff Beam (UNC Couch Botanical
Library) for help in obtaining literature, and, especially, to Susan Whitfield
(Dept. of Biology, UNC-CH) for the fine illustration.

REFERENCES

ARRIAGADA, J.E. 1997. The genera of ae eae (Compositae; Asteraceae) in the southeastern

Jnited States. Harvard Pa 1p. Bor —48
Cabrera, A.L. 1961. Obser “obs I a Inuleae — Gnaphalinae (Compositae) de América
del Sur. Bol. Soc. A paahie. o6e 9: 359-386.

CABRERA, e L. 1963. Flora de la Provincia de Buenos Aires. Parte VI. Compuestas. Colecci6n
rientifica del I.N.T.A. Buenos Aires, Argentina.
Drury, D.G. 1971. The American spicate oan 1s adventive to New Zealand: (Gnaphalinm
section eee one Compositae). New Zealand J. Bor. 9:157—185.

GODFREY, . A synopsis of Cae (Compositae) in the southeastern United
States. Guan. i . rida Acad. Sci. 21:177-184.
Nesom, G.L. The taxonomic status of Gamochaeta (Asteraceae: Inuleae) and the spe-

cies of the nite States. Phytologia 68:186—-198
Pruskl, J.F. 1997. Asteraceae. Flora of the Genemuelan aries 3:177-393.
Webs, C.J. 1988. Gnaphalinm. Flora of New Zealand 4:234—249

VICIA LUTEA (FABACEAE) NEW TO TEXAS

AMANDA K. NEILL
Department of Biology Herbarium
Texas AGM University
College Station, TX 77843-3258, U.S.A.

netlla@ mail. bio.tamu.edu

Vicia lutea L. (yellow vetch) has not been reported for Texas by Correll and
Johnston (1970), Hatch et al. (1990), Jones et al. (1997), or Diggs et al.
(1999). This species is easily distinguished from the other vetches found in
Texas by its axillary sessile flowers, large yellow corollas, unequal calyx lobes,
and pustulate-pilose fruits. Isely (1998) provides a key to separate this Vicia
from others in the United States.

Vicia lutea is a weedy annual native to the Mediterranean region. How-
ever, the species is known to occur sporadically in North Carolina and Louisiana,
and is seemingly naturalized in California (Hickman 1993). Thomas and
Allen (1998) mapped it in nine northern Louisiana parishes. Personal com-
munications with other herbaria have revealed additional collections:

MA. Lee Co.: 11 May 1961, Kendrick 83 (AUA). Sumter Co.: | May 1994,
Thomas & Thomas 135, 206 (BRIT). OREGON. Lane Co.: 17 Jun 1998, Koenig s.n. (OSC).

During field work for a county flora in east central Texas, Vicia lutea was
collected in Madison County. A single, dense population of Vicia lutea was
found in full anthesis along seven meters of a dry, sandy slope. The slope
was along a roadside, in full sun, and adjacent to a gravel pit. Two weeks
later the plants were leafless, yet held a large amount of ripening fruit. Other
herbaceous species on the site included Polygala polygama, Coreopsis basalis,
Lathyrus hirsutus, Helianthemum georgianum, Schizachyrinm scopavium, and Paspalum
plicatulum. Woody vegetation along the nearby fenceline was composed of
Quercus marilandica, Q. falcata, and Pinus taeda.

The plants at the Madison County site were found at a relatively undis-
turbed roadside area that is not mowed. The presence of this population
may be attributed to traffic associated with a nearby gravel pit, or to cattle
and hay transport through this rural area. Upon inspection of previous Madison
County collections not yet identified, another specimen of Vicia lutea was
found. That individual [Madison Co.: 11 April 1998. A.K. Ne// 1458 (TAMU)}
was collected at anthesis approximately 27 miles southwest of the voucher
specimen, but the reproductive success of that population has not been es-
tablished.

Vicia lutea may remain a waif in humid, non-Mediterranean climates. One

Sipa 18(4): 1265-1266. 1999

1266 Sipa 18(4)

visible effect of humidity on the Madison County population was an infec-
tion with powdery mildew fungus; the mycelial layer imparted a silvery
cast to the leaf surfaces. Additional collections of this species may be ex-
pected from Texas and other warm-climate states. Images of the voucher
specimen are available at Texas A&M’s Bioinformatics Working Group Image
Gallery page (http://www.csdl.tamu.edu/FLOR A/ imaxxfab. htm)

Voucher specimen: U.S.A. TEXAS. acs Co.: Roadside cut-bank of Chazos loamy
sand at entrance to gravel pit, W side of FM 247, 1.8 miS of TX Hwy 21 jcc. (at els
Sand Ridge Quadrant, as 217 ft, 30°00'N, ee "W, 1 May 1999, A.K. Nei// 2618 (TAMU

ACKNOWLEDGMENTS

Iam grateful to the following curators and institutions who supplied
collection data: Bonnie Amos (SAT), Anne Bradburn (NO), Steve Broich
(OSC), Larry Brown (SBSC), Richard Halse (OSC), Robert Haynes (UNA),
Leigh Johnson (NCSC), Jim Massey (UNC), Mark Mayfield (LSU), Scott
Sundburg (OSC), Jerome Ward (AUA), Tom Wendt (LL, TEX), Donna Ford-
Werntz (W VA), Robert Wilbur (DUKE), and Lindsay Woodruff (BRIT).
Monique Reed (TAMU) provided assistance in identifying this species, and
Hugh Wilson, my advisor, gave support and encouragement during my thesis
work in Madison County.

REFERENCES

Corrett, D.S. and M.C. Jounsron. 1970. Manual of the vascular plants of Texas. Texas
Rese a Foundation, Renner

Diccs, G.M., B.L. Lipscomp, and R.J. O'KENNON. 1999, Shinners and Mahler's illustrated
flora of ce 1 Central Texas. Sida, Bot. Mis

Harcu, S.L., K.N. GaNbru, and L.E. a. A ). Checklist oft the vascular plants of Texas.
Texas Agric. | Sta. Bull. MP-1655

Hickman, J.C. (Ed.). 1993. The Jepson sete: Higher plants of California. Univ. of Cali-
fornia Press, oak ley

Isety, D. 1998. Native oot naturalized Leguminosae (Fabaceae) of the United States (ex-
clusive of Alaska and Hawaii). Monte L. Bean Life Science Museum, Brigham Young

Iniv., Provo, UT.

Jones, $.D., J.K. Wiper, and P.M. Monrcomery. 1997. Vascular plants of Texas, a compre-
hensive checklist including synonymy, bibliography, and index. Univ. of Texas Press,
Austin.

THomas, R.D. and C.M. ALLEN. 1998. Atlas of the vascular flora of Louisiana. Vol. III: Dicotyledons:
Fabaceae-Zygophyllaceae. Louisiana Dept. of Wildlife and Fisheries, Baton Rouge.

IPOMOEA ASARIFOLIA (CONVOLVULACEAE),
ANOTHER POTENTIAL EXOTIC PEST
IN THE UNITED STATES

KATHLEEN CRADDOCK BURKS
Florida Department of Environmental Protection
3915 Commonwealth Blrd.,
Tallahassee, FL 32399, U.S.A.
kathy. burks@ dep. state.fl.us
DANIEL FE. AUSTIN

Florida Atlantic University
777 Glades Road
Boca Raton, FL 33431, U.S.A.
daustin@ fau.edu

Known 1n Brazil as “salsa,” [pomoea asarifolia (Desr.) Roem. & Schult. is currently
distributed in both the New and Old World but is of uncertain origin; re-
cent research suggests Asia as the most likely native region (Austin, unpubl.
data). In 1994, a population of this species was discovered in Broward County,
Florida, the first known occurrence in the United States. Based on herbarium
specimens collected at the time and identified by the second author, the
species was subsequently recorded by Wunderlin (1998). In this note, we
wish to formally report details of the discovery and provide additional in-
formation on the species’ characters and habits, which have raised concerns
about its potential as a pest plant in this country.

Following up on a report to the first author from a wetlands botanist
inspecting a delineation project, aquatic regional biologist Jackie Jordan)
Smith investigated in October 1994 a suspected population of Ipomoea aquatica
Forssk., a prohibited species in Florida (Florida Administrative Code, DEP
62C-52). Though the suspect vine was found growing across the surface of
a pond, as [. aguatica commonly grows, it was not that species and did not
match any other /pomoea known for Florida. A small sample was identified
in November 1994 by the second author as a species new to the United
States. In August 1996, the site, located along a canal maintained by the
South Broward Drainage District (SBDD), was revisited, and fresh flowering
specimens were collected for verification. The vine was well-established around
and in a small pond at a pasture edge and clambered up adjacent shrubs
(Myrica cerifera L.); it also appeared to have spread farther by runners among
the mowed grasses down and along the canal bank.

Sipa 18(4): 1267-1272. 1999

1268 Sipa 18(4)

The site has been dramatically altered since 1996 by urban development
activities. By June 1999, much of the SBDD Canal No. 5 had been filled in
and paved over with the opening of the divided, four-laned Dykes Road. A
large residential subdivision, Silver Shores, occupies much of the former
pastureland just north of the earlier-visited small pond, a remnant of which
still exists in the remaining low, wet area, which itself is slated for residen-
tial development and has experienced some preliminary bulldozer work.
The I. asarifolia survived this massive disturbance; in fact, the population
has spread considerably along the new roadway and away from the pond
area, creeping across bare lime rock, floating across shallow pools of water,
and clambering over native and exotic herbs and shrubs. The Sopulstion
extends about 800 m along Dykes Road and as far as 300 m westward from
it, between Silver Shores and Miramar Parkway. It does not appear to have
been carried east of Dykes Road or south of Miramar Parkway, areas also
undergoing considerable earth-moving activity.

Vouchet Sees U.S.A. FLORIDA. Broward Co.: W side right-of-way, SBDD Canal
No. 5, just NW of I-7 pan Miramar Parkway interchange, within oe limits, in

mowed area and around small adjacent pond, on sandy soil, 17 Oct i, Jackie Jordan s.n.
(FAU, FSU); 21 Augusce 1996, Burks 1074 (FAU, FLAS, FSU, USE); oe W side of Dykes

Rd., in disturbed wet ground, thin muck or sand over limestone or bare limestone, to 300

a

m W of roadside, between southern boundary of Silver Shores subdivision and ject. of Dykes
Rd. and Miramar Parkway, just NW of I-75 and Miramar Parkway interchange, within
Miramar city limits, SE 1/4 of NE 1/4 Sec 29, TSIS R40E, Lat. 25° 59.158 N, Long. 80°
21.669 W, 25 Jun 1999, Burks 1159 (FAU, FLAS, FSU, USF).

A vector for this introduction is unknown, but the possibilities are nu-
merous. The site has a long history of disturbance: cattle pasture, canal bank,
roadside, housing development; the stand first noted in 1994 may have been
overlooked for some time. Seed or stem fragments could have arrived on
farm or canal-maintenance equipment; as a contaminant in feed, straw, or
grass seed; or in the gut of imported cattle (or migratory birds—little is
known about the consumption/dispersal of 1. ee by animals). Dumping
of imported packing material or horticultural material (yard trash and white
goods”) was also a common practice in the area. The species is not known
in cultivation in the United States, but is cultivated for ornament in Bra-
zil, as “salsa” or “salsa-brava” (Lorenzi & Moreira de Souza 1999). Yet an-
other possibility is introduction for use in traditional herbal remedies (Austin,
unpubl. data).

Although 1. asarifolia may not be New World tn origin, it is found widely
in Tropical America, from the Carribean to Paraguay (Adams 1972; Austin

1975, 1982a-c, 1997, 1998a-b; Austin & Cavalcante 1982; Austin & Huaman
1996; Austin & Staples 1981; Leon & Alain 1974; McDonald 1994). It is
ilso known in Africa (Heine 1963; Rendle 1905), and in Asia, from Bali,

BuRKS AND AUSTIN, Ipomoea asarifolia 1269

East Java, India, and West Pakistan (Austin & Ghazanfar 1979; Matthew
1995; Ooststroom 1953).

Documented habitats for this perennial species are largely wet sites—
marshes, swamps, roadside ditches, and in wet-cultivated crops such as rice
(Austin 1982a). While little has been reported in the literature about J.
asavifolia as a pest plant across its range, it isa recognized agricultural weed
in Venezuela and Brazil (Austin 1982c, 1998a; Kissman & Groth 1992;
Lorenzi 1991; Moacyr et al. 1995). The plant is sensitive to frost, but grows
in a wide range of moist soils and is easily propagated by stem fragments or
seeds (Lorenzi & Moreira de Souza 1999). It apparently produces viable seed
in Florida; a few seedlings were observed at the Miramar site, along with
extensive mats of vines freely rooting at the nodes. Those vines seen in 1999
on drier ground and mixed with other herbaceous vegetation [e.g., Setaria
parviflora (Poir.) Kerguélen, Expatorium capillifolium (Lam.) Small} often had
leaves noticeably damaged by insects, while vines growing on more open,
saturated ground showed no damage. A 1996 sample of unrooted vine was
floated in a vat of well water in a quarantine greenhouse (Tallahassee, Florida);
it sprouted roots at the nodes and sustained itself under those conditions
for several months, until it was moved to a frequently watered pot of soil,
where it still grows.

The species has been placed taxonomically in the same section of Ipomoea
[subg. Eriospermum (Hall f.) Verdc. ex D.F. Austin sect. Erpipomoea Choisy]
as 1, aquatica and I. pes-caprae (L.) R. Br., species similarly known for their
proclivity to spread vegetatively by oo adventitiously rooting, runners.
1. asavifolia resembles I. pes-caprae in habit, flower color, and general leaf
form; however, the former is a species of freshwater habitats whereas the
latter is confined to saline conditions near seacoasts. Also, I. asarifolia never
has the emarginate to bilobed leaf apex characteristic of 1. pes-caprae.

Diagnostic characters for identifying I. asarifolia in the field include its
habit of horizontal stems and alternate, smooth, dark-green, cordate leaves,
these often with purplish venation and with tips obtuse to mucronulate.
The stems, whether clambering or horizontal, may have small, sparse, fleshy
trichomes. The plant's showy morning-glory flowers have a campanulate to
funnelform corolla of deep rose-lavender with a purple throat (Fig. 1).

Given the spreading habit of this species, its apparent preference for wetlands,
its adaptability to disturbance, its recognition as a weed in some locales,
and our experience with the Florida population, we view this introduction
with alarm and will continue to monitor its occurrence. Steps are immedi-
ately being taken to confine any excavated fill to the current site and to

plan a control effort
We gratefully acknowledge John Tobe for noticing the suspect morning

—_

1270 Sipa 18(4

Fic. 1. [pomoea asarifolia in Florida. Top: Flower, leaf form; Bottom: Creeping habit over
water, bare ground, other plants.

BURKS AND AUSTIN, Ipomoea asarifolia 1271

glory and Jackie Smith for initially investigating the report, and provid-
ing, along with Joe Certain of SBDD, assistance in the field. We also thank
Loran Anderson for helpful comments on an earlier draft of this note.

REFERENCES

Apams, C.D. 1972. Flowering plants of Jamaica. Univ. of the West Indies, Mona.
Austin, D.F. 1975. Convolvulaceae. In: Flora of Panama. Ann. Missouri Bot. Gard, 62:147—
224.

Se Matters of meaning: salsa cea Bull. Fairchild Trop. Gard. 37:24—30.
—_____. 1982b. Convolvulaceae, Fam. 165. In: G. oe and B. Sparre, eds. Flora
of Ecuador Ne = Swedish Research Council Rae
—__.. 1982¢. Convolvulaceae. In: Z. Luces de om and J.A. Steyermark, eds.
Flora de Venenele. Vol. 8, pt. 3. Ediciones Fundacié6n, Educacié6n Ambiental, Caracas.
Pp. 15-226.
—____.. 1997. Convolvulaceae. In: Boggan, J., V. Funk, C. Kelloff, M. Hoff, G. Cremers,
and C. Feuillet, eds. Checklist of the plants of the Canc (Guyana, Surinam, French
Giana: 2nd Ed. Publication 30, Biological Diversity of the Guianas Program, Dept.

of Botany, oo Museum of Natural History, Smithsonian Institution, Washington,
DC. Pp. 8

ps ee Convolvulaceae. In: Steyermark, J. A., P. E. Berry and B. K. Holst

eds. Flora of the Venezuelan Guyana. Volume 4. Caesalpinaceae-Ericaceae. Missouri Bot.
Gard. Press, St. Louis. Pp. 377-424.

—____. 1998b. Convolvulaceae. In: Cremers, G. and M. Hoff. Inventaire taxonomique
des plantes de la Guyane Francaise VII—-Les Dicotyledones. 3éme Partie: Connaraceae a
Fabaceae. Muséum National d'Histoire Naturelle, aes d’Ecologie et de Gestion d la
Biodiversité, Ministére de l'Environnement, Paris. Pp. 18—32.

Austin, D.F. and P.B. Cavalcante. — Convolv Hee de Amazénia. Publ. Avulsas No.
36. Museu Emilio Goeldi, B

Austin, D.F. and S. ee oo Convolvulaceae. In: E. Nasir and A.J. Ali, eds. Flora
of West Pakistan. Vol. 126. Agricultural Research Council Islamabad. Pp. 1-64

Austin, D.F. and Z. JINAMAN. 1996. A synopsis of Ipomoea (Convolvulaceae) in the Ameri-
cas. Taxon 45:3

Austin, D.F. and G. W. STaPLes. 1981. Convolvulaceae. In: B. Maguire, ed. Botany of the
Guayana Highlands. Mem. N.Y. Bot. Gard. 32: 309-323.

HEINE, H. (ed.). 1963. Convolvulaceae. In: Hutchinson, J. and ee. _ M. (revised by

RW. Keay and EN. sae Flora of West Tropical Africa. Vol. ondon, Crown
Agents (HMSO). Pp. 337-
See K.G. and D. Grortn. ioe. Plantas infestantes e nocivas. Tomo II. BASF Brasileira
A., Sao Paulo, Brazil.

ta H. and H. Aan. 1974. Flora de Cuba. Vol. 2. Otto Koeltz Science fee hers, FOROS

Lorenzi, H. 1991. Plantas daninhas do Brasil: Terrestres, aquaticas, parasita
Weeds of Brazil. 2a ed. Editora Plantarum, Nova Odessa, SP.

Lorenzi, H. and H. Moreira br Souza. 1999. Plantas ornamentais no 0 Brasil arbustivas,
herbaceas e trepadeiras. 2a ed. Instituto Plantarum, Nova Odessa

MatruHew, M.H. 1995. An excursion flora of Central Tamilnadu, India. i A. Bales Uitgevers
b.v., Rotterdam, The Netherlands.

Moacyr, B.D., J.A. Wisr, and T.E. Dawson. 1995. Root and shoot growth in response to
soil drying in four Amazonian weed species. Revista Brasileira de Fisiologia Vegetal 7:53—
a2

cae

1272 Sipa 18(4)
McDonatp, A. 1994. Flora de Veracruz. Convolvulaceae H. Instituto de Ecologia, A.C.,
Xalapa, Veracruz, and Univ. of California, Riverside.
OosrsTRooM, S.J. VAN. 1953. Convolvulaceae. Flora Malesiana. Djakarta, Noordhoof-Kolf!

Pp. 388-512.

RENbLE, A.B. 1905. Convolvulaceae. In: Thiselton-Dyer, W.T., ed. Flora of tropical Africa. L.
Reeve & Co., Kent, England. reprint by Beadley Brothers Ltd., Kent, 1959. Vol. 4(2):62—206,

WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of
Florida, Gainesville, FL.

PAEDERIA FOETIDA (RUBIACEAE),
NEW TO THE FLORA OF NORTH CAROLINA

PETE DIAMOND

Department of Horticulture
North Carolina Zoological Park
Asheboro, NC 27203, U.S.A.

During the summer of 1998, Paederia foetida L. (Rubiaceae) (Fig. 1) was
discovered growing at the North Carolina Zoological Park, five miles south
of Asheboro, North Carolina, in Randolph County. The plant apparently is
naturalized in its location from an unidentifiable source. The native veg-
etation around the collection location, adjacent to a zoological exhibit building,
is composed of hardwood, deciduous forest species. The landscape is supplemented
with additional plantings of native species to highlight the state’s piedmont

and mountain geographic regions.

Voucher specimens: NORTH CAROLINA. fea a a North <n Zoologi-
cal Park, Asheboro, ase 37'52.37934" N, 79° 36.1 “WY, 13 Aug 1998, Diamond
28 (BRIT, NCU, NC Zoological sere North - arolin né on -al Park, peace 35°
> 52.37934" N, 79° 45’ 36.14607" W, 19 Oct 1998, oe (NCU, NC Zoologi-

cal Park).

While often encountered in horticultural reference materials and select
floras as Paederia scandens (Lour.) Merr. (Hillier Nurseries 1991; Kriissman
1977; Walker 1976), P. foetida L. is currently recognized as the correct scientific
name (Kartesz 1999). Synonymous names include P. chinensis Hance and P.
tomentosa Maxim. (Kartesz 199

Paederia foetida is a twining, deciduous vine capable of nodal rooting and
notable for the rank odor it releases on warm, humid days and when parts
of the plant are bruised. In Japan, the plant is referred to as “Flatulent Vine”
(Walker 1976), while in the U.S. it has earned the name “Skunk-vine” (Weakley
1998). The genus comprises about 20 species of climbing shrubs native to
temperate and tropical Asia and South America. Leaves of P. /oetida are ovate
to broadly ovate, truncate, cordate or subcordate, opposite, the veins on both
surfaces with appressed pubescence, the upper surface also bearing short
hispid or papillose-hispid hairs. Panicles are axillary; calyx lobes < 1 mm
ong at flowering; corolla white, tubular, cc. 1 cm long, with deep red throat.
Fruit orange, globose, a 2-locular berry.

Documentation of the occurrence of P. fuetida in North America is lim-
ited. Radford et al. (1964) noted it was “collected spreading from its site of
cultivation in Darlington, Co., SC,” after the manuscript for the Manual of

SIDA 18(4): 1273-1276. 1999

Stipa 18(4)

Herbarium of the
North Carolina Zoological Park
aederia ida
Aid
long, white, granular
in-anilary cymes
}
FIG

I 4 Corolla tubular
throat dark red, styles 2
leaf surf: b
Pl

Jem
2. Flowers 5-merous.
Il il
|
{
Streamside building
Zoologice

Stems twining or
becoming 1

gical Park, Asheboro, NC. unknown
coll. Pete Diamond, no 428

Paederia foetida. North Carolina Zoological Park, Asheboro, Diamond 479

3 August 1998

DIAMOND, Paederia foetida 1275

the Vascular Flora of the Carolinas had gone to press. There is little docu-
mented change in the distribution of P. foetida in the Carolinas during the
past 30 years. Conferring with Radford, Weakley (1998) reports P. foetida
is limited to disturbed areas of the South Carolina coastal plain and rarely
spreads from plantings. Nelson (personal communication, 1998) notes that
P. foetida is fairly widespread and naturalized in warmer, Gulf coastal states
of the U.S., notably Florida. There also are documented reports of P. foetida
in Louisiana (Thomas & Allen 1997) and Texas (Brown 1998). The species
was first recorded as naturalized from O’ahu, Hawaii, in 1854 (Wagner et
al. 1990).

Among three varieties of P. foetida recognized by Walker (1976, as P. scandens),
the North Carolina plants are apparently closest to P. scandens var. mairei
(Lév'l) Hara, based on the leaf shape and vestiture. In var. mairei, Walker
noted that “lower leaf surfaces are glabrous except for tufts of hairs in vein
axils,” while in var. vz//osa lower leaf surfaces are villous. Leaves of the North
Carolina plants are sparsely hairy but not distinctly villous.

A record of Paederia foetida in North Carolina is noteworthy not only to
document the spread of this exotic species but also to confirm its ability to
tolerate severe winter temperatures. In North Carolina the plant is prob-
ably near the limit of its northern hardiness range. This species is report-
edly capable of surviving minimum winter temperatures of 0 to 10 degrees
Fahrenheit (Huxley 1992), although Kriissman (1977) suggests a cold har-
diness of —10 to 0 degrees Fahrenheit.

Paederia foetida has proved hardy in North Carolina, surviving as an her-
baceous perennial through the winter of 1998/1999 and, in all likelihood,
several previous winters as well. Efforts were made in October 1998 to limit
the spread of this species by pulling out or cutting back stems, largely to
keep the plant from establishing itselfand becoming invasive. Pruned stems,
however, were resprouting and leafing out by mid-April 1999.

The single plant discovered at the North Carolina Zoological Park was
extensive, climbing through nearby shrubs and low branches of trees and
scrambling over the ground up to four meters in length. Because of the propensity
for nodal rooting, the main stem is difficult to determine. The most likely
point of origin, however, appears near the trunk of a beech tree (Fagus grandifolid),
This tree was planted during early landscaping around the building in December
1993. Deliveries of 14 balled and burlapped FE. grandifolia were received at
the Park in mid-November from two separate nurseries in the vicinity of
McMinnville, Tennessee. While there are currently no reports of P. foetida
occurring in Tennessee, one possibility is that Paederia was already grow-
ing in the root ball of one of these trees. The plant may also have sprouted
from a seed deposited by migratory birds or, possibly, from purchased com-

1276 Stipa 18(4)

mercial birdseed of unknown origin that was scattered in the vicinity. Be-
cause the plant was growing at the Park for an undetermined amount of
time, any theories relating to its original source are inconclusive.

Several fruits were observed in October 1998, but reproduction appears
low. Herbarium specimens collected from the NC Zoological Park indicate
that only two berries were produced from more than two dozen panicles,
suggesting the occurrence of a low frequency of successful self-pollination.
By contrast, fruiting specimens (NCU) collected in Japan in 1985, show
heavy berry production.

REFERENCES
aie N, L. 1998. Personal communication, Houston Community College and Herbarium
SBSC, Houston (via Biota of North America Program, NCNB, UNC, Chapel Hill, NC).
ata seen (COMPILER). LOOL. The Hil 3 manual of trees and shrubs. Sixth edition.
David and harles, Ltd., Newton Abbor, U.K.

Karresz, J.T. 1999. A synonymized cee and atlas with paeee attributes for the
vascular ora of the United States, Canada, and Greenland. First Edition. In: Kartesz,
J.T. and C.A. Meacham. Synthesis of A Norch American Flora, soe 1.0. North Carolina

Botanical Garden, Chapel Hill,

KrussMAN, G. 1977. Manual of se broad-leaved trees and shrubs. Volume 2. Tim-
ber Press, oo and OR.

Netson, J. 1998. Personal communication, Unity. of South Carolina at Columbia herbarium.

Hux.ey, A. (ed. in chief). 1992. New Royal Horticultural Society dictionary of gardening.
Volume 3. MacMillan Press Led., London.

Raprorb, A.E., H.E. Antes, and C.R. Brett. 1964, ‘anil of the vascular flora of the Caro-
linas. Univ. oe North Carolina Press, Chapel Hil

Tuomas, R.D. and C.M. ALLEN. 1997. Atlas of the ae flora of Louisiana. Volume 3.
Louisiana Department of Wildlife and Fisheries, Natural Heritage Program, Baton Rouge.
Wacner, W.L., D.R. Herpst, and $.H. Somer. 1990. Manual of the flowering plants of

Hawaii. Univ. of Hawaii Press, Bishop Museum, Honolulu
a KER, E.H. 1976. Flora of Okinawa and the Southern Ryukyu Is! ce Smithsonian Institution
ress, ae C
WeraAkLEY, A. 1998. Flora of the Carolinas and Virginia, wor

king draft.

LAPSANA COMMUNIS (ASTERACEAE)
NEW FOR TEXAS AND NOTES ON INVASIVE
EXOTICS

ROBERT J. O.KENNON,! THEODORE M. BARKLEY;,!
GEORGE M. DIGGS, JR.,':? BARNEY LIPSCOMB!

‘Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102, U.S.A.
?Department of Biology
Austin College
Sherman, TX 75090, U.S.A.

okennon@brit.org, barkley@brit.org
ediges @austinc.edu, barney@brit.org

Lapsana communis L., nipplewort or succory dock-cress, is a native of Europe
(Vuilleumier 1973; Sell 1976; Stebbins 1993) naturalized in a variety of
localities in the United States. It is usually described as a weed inhabiting
roadsides, disturbed places, and waste areas, often in shady situations. A
genus of nine species, Lapsana (Asteraceae: Lactuceae) is native to the ex-
tratropical regions of Europe, Asia, and northern Africa. The only truly weedy
member of the genus, L. communis is also adventive in at least Africa, Polynesia,
the West Indies, and South America (Vuilleumier 1973). In North America
the species is known from Ontario, Quebec, and the northeast U.S. south
to North Carolina and west to North Dakota, Missouri and Arkansas; it is
also found from British Columbia south to Oregon and California (Robinson
& Fernald 1908; Britton & Brown 1898, 1913; Fernald 1950; Gleason 1952;
Gleason & Cronquist 1963, 1991; Steyermark 1963; Radford et al. 1968;
Strausbaugh & Core 1978; Cronquist 1980; Barkley 1986; Smith 1988;
Stebbins 1993). Voss (1996), for example, cited the species for Michigan,
indicating it had been found sparingly in the past and largely overlooked
until very recently, with it now being very abundant in some localities. Lapsana
communis was not reported from Oklahoma (Taylor & Taylor 1994); how-
ever, it was reported from three western counties of Arkansas (Smith 1988).
Despite this proximity, the species has not been previously reported from
Texas (Correll & Johnston 1970; Stanford 1976; Hatch et al. 1990; Jones
et al. 1997; Diggs et al. 1999).

A collection made in 1999 in Fort Worth (Tarrant County) is apparently
the first documented occurrence of this species for Texas. The identification

Sipa 18(4): 1277-1283. 1999

1278 Sipa 18(4)

was determined by Theodore M. Barkley of BRIT (formerly, Kansas State
University).

Fort

Voucher specimens: TEXAS. Tarrant Co.: ey area in landscape, 301 Crestwood,
Worth, 25 Jun 1999, i ’"Kennon 14377 (BRIT
At the collection locality a single individual was observed as a landscape
weed. The plant was 70 cm tall and 25 cm wide. This yard has incurred a
large number of weedy species after a truckload of compost, from a large
compost operation run by a local nursery, was dumped and spread in 1997.
Seeds of various weedy species, including Lactuca saligna (O’Kennon et al.
L998a), Plantago coronopus (O’Kennon et al. 1998b), and Lapsana communis,
were apparently introduced in this manner. Other populations of Lapsana
have not been found in North Central Texas, and it is unclear whether the
species will gia more widely established. However, it would not be surprising
that the species has been spread to other landscapes by the same method.
The generic name Lapsana is derived from Greek, /apsane or lampsane, a
name given by Dioscorides to a vegetable, apparently a species of Raphanus
(Vuilleumier 1973). According to Vuilleumier (1973), Bentham and Hooker
(1873) allied Lapsana with A pogon ELL. (now included in Krigza L.), but Stebbins
(1953) considered Lapsana (because of its lack of a pappus) to be a special-
ized offshoot of the Youngza-Ixeris line of the Crepidinae. Jeffrey (1966) in-
dicated a similar relationship, putting Lapsana in his “Crepis-series” with
Crepis, Ixerts, and Youngia. Small (1917, cited in Vuilleumier 1973) reported
that when stamens of this species are touched, the filament nearest the stimulus

—

contracts and the entire anther tube faces the visitor as the style pushes out
the pollen toward it. Vuilleumier (1973) noted that the milky juice of the
plant is soothing to sensitive skin, particularly on the nipples of nursing
mothers (hence the common name nipplewort).

Lapsana communis (Fig. 1), the flowers of which resemble a Lactuca, Crepis,
Hreracium, or Youngia, can be recognized by the following description (modified
from Cronquist 1980 and Barkley 1986); the absence of a pappus is par-
ticularly noteworthy. Erect, branching to single-stemmed annual herb [acting
as a biennial in our area}, 1.5—10(-15) dm tall, hirsute to glabrescent, wit

—
—
~

milky juice; leaves alternate, petiolate, chin, ovate to subround, obtuse to
rounded, variously dentate, or the lowermost lyrate [somewhat resembling
basal leaves of members of the mustard family}, 2.5—10 cm long, to 7 cm
wide, progressively shorter-petiolate upward, the uppermost blades distinctly
narrowed; heads few—numerous in a corymb-like to somewhat elongate panicle-

like capitulescence; involucre 5-8 mm tall, cylindric to campanulate-spreading;
principal phyllaries ca 8, subequal, uniseriate, subtended by minute caly-
culate bracts; receptacle naked; florets 8-15 per head, all ligulate and fer-

tile, corollas yellow; achenes 3-5 mm long, + terete to weakly flattened,

O'KENNON ET AL., Lapsana communis 1279

aa glabrous, with numerous nerves (ca. 18-30); pappus none; (7 = 6,
7, 8). Jun—Sep. The accompanying illustration is reprinted from Gleason
(1952) and Strausbaugh and Core (1978) with the permission of the pub-
lishers.
NOTES ON INVASIVE EXOTICS

Lapsana communis is one of a continuing string of introduced species (defined
here as those originating outside the United States) introduced into the flora
of Texas. These taxa are variously referred to as alien, exotic, or foreign. In
a recent floristic study on North Central Texas (Diggs et al. 1999), it was
determined that 17.7% (or 394 species) of the flora consisted of such intro-
duced species. This is roughly what would be expected based on data from
other parts of the United States. For example, the level of exotic species in
the northeastern states has been estimated at 22% (Elias 1977) and 20—
30% (Stuckey & Barkley (1993). Comparable figures for introduced spe-
cies for other states include California (17.5%) (Rejmanek & Randall 1994),
Colorado (16%), Iowa (22.3%), Kansas (17.4%), and North Dakota (15%)
(Stuckey & Barkley 1993). Considering how recently North Central Texas
was settled and converted from almost totally native habitat, the 17.7%
figure is quite striking.

Some of these exotics are extremely invasive taxa capable of becoming
serious agricultural pests or of destroying native habitats. Particularly problematic
are those that aggressively invade native ecosystems, reproduce extensively,
and occupy the small remaining areas of natural habitat used by indigenous
species. A few of the many problematic species currently spreading in North
Central Texas include Carduus nutans L. subsp. macrocephalus (Desf.) Nyman
(musk-thistle or nodding-thistle), Hydrilla verticillata (L.f.) Royle (water-
thyme), Pweraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & Almeida
(kudzu), and Scabiosa atropurpurea L. (pincushions, sweet scabious). Scabiosa,
for example, is now known from at least seven counties (Collin, Dallas, Fannin,
Grayson, Hopkins, Hunt, and Lamar) and will almost surely be found in
many more. In Collin County, it lines some roadsides (e.g., Hwy. 75) and
virtually forms monocultures. Given the ease with which it moves into adjacent
fields, there is real concern that sweet scabious may be able to invade native
prairie remnants. Likewise, native habitats (e.g., forests, prairies) are being
invaded by a host of other exotics, including Lonicera japonica Thunb. (Japanese
honeysuckle), Liguwstrim species (privets), Sorghum halapense (L.) Pers. Johnson
grass), and Festuca arundinacea Schreb., (tall fescue).

In many areas, a high percentage of the plants encountered will be of
introduced species. Given the speed with which exotics have invaded, and
the percentage of habitat they now occupy, after habitat destruction, inva-

1280 Sipa 18(4)

Fic. 1. Habit of Lapsana communis (Britton & Brown 1913, with permission, New York
Botanical Garden, and Strausbaugh and Core (1978) with permission Seneca Books, Inc.).

sion by exotics may be the most serious threat facing native plants in North
Central Texas (Diggs et al. 1999). Problematically, this 1s not a short-term
concern. According to Cronk and Fuller (1995), the introduction of exotics
is “...a lasting threat because when exploitation or pollution stops, ecosys-
tems often begin to recover. However, when the introduction of alien or-
ganisms stops the existing aliens do not disappear; in contrast they some-
times continue to spread and consolidate, and so may be called a more pervasive
threat.” Unfortunately, there is a continuing lack of recognition of the problem.
For example, the Federal noxious weed Sa/vinia molesta Mitchell has recently
been found in abundance at Toledo Bend Reservoir, on the Texas-Louisiana
border (Jacono 1999). This native of Brazil (Forno 1983) can grow rapidly
and cover the surface of lakes and streams, and the floating mats shade and
crowd out native plants. Additionally, the thick mats reduce oxygen con-
tent, degrade water quality, and can cause physical problems including hindering
boats and clogging water intakes (Jacono 1998). In Texas, it is considered
a “harmful or potentially harmful exotic plant” ae it is illegal to release,
import, sell, purchase, propagate, or possess this species in the state (Harvey
1998). Nonetheless, likely sources for the introduction are local nurseries

O'KENNON ET AL., Lapsana communis 1281

found distributing the species as an ornamental water garden plant (Randy
Helton, pers. comm., in Jacono 1999).

nother example, Lythrum salicaria L. (purple loosestrife), can also be found
for sale in Texas nurseries and is even planted on local college campuses. In
some parts of the northeastern United States, L. salicaria aggressively in-
vades native marshlands eliminating native species. Dense stands covering
thousands of acres are sometimes formed with even tenacious natives such
as Typha (cat-tail) species being excluded. Purple loosestrife is often cited
as one of the most detrimental cases of habitat alteration by an exotic spe-
cies in the U.S. It was introduced in New England in the early 1800s and
by 1995 was known in every state but Florida. Because of its potential as a
pest, it has been declared a noxious weed in several states with laws ban-
ning its distribution and cultivation; this species should not be planted (Stuckey
1980; Graham 1986; Yatskievych & Spellenberg 1993; Flack & Furlow 1996);
htcp://plants.usda.gov/.

A final example for Texas is Sapimm sebiferum (L.) Roxb., (Chinese tallow
tree, vegetable tallow tree). This rapidly growing tree was introduced into
the U.S. in South Carolina in the late 1770s and is now widespread; it dis-
places native vegetation and is considered one of the most serious invasive
exotics in the U.S. It apparently releases compounds that modify soil chemistry
and affect the establishment of native species (Flack & Furlow 1996). In
Texas, it is extremely problematic in invading areas of native coastal prairie
and East Texas forest (e.g., Big Thicket National Preserve). While the seri-
ousness of the threat of exotics is still significantly underrealized, at least
some increase in awareness is occurring. Examples of recent articles in the
popular press on exotics include one in the Nature Conservancy Magazine (Flack
& Furlow 1996) and a just published report in Scientific American discuss-
ing the serious invasion of Lygodinm microphyullum (Cavanilles) R. Brown
(Old World climbing fern) in Florida and the threat it poses to the Ever-
glades (Mirsky 1999).

REFERENCES
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Univ. Press of Kansas, Lawrence. Pp. 838-102
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2500).

Britron, N.L. and A. Brown. 1898. An illustrated flora of the northern United States,

Canada and the British ame Charles Scribner’s Sons, New York.
1913. An illustrated flora of the northern United States,

Canada and = British possessions. "New York Botanical Garden, Bronx.

CorreLl, D.S. and M.C. JoHNsron. 1970. Manual of the vascular lanes of Texas. Texas
Research Foundation, Renne

Cronk, Q.C.B. and J.L. pune r. 1995. Plant invaders: the threat to natural ecosystems.
Chapman ue Hall, New York.

1282 Sipa 1L8(4

wa

CRONQUIST, L980. Asteraceae. Vascular flora of the southeastern United States 1:1—26 1.
IV, | North Carolina Press, Chapel Hill.
Diccs, G M. Jr., BLL. Lipscoma, and R.J. O'RENNON. 1999. Shinners & Mahler's illuscraced
flora of nee seus ee cas. deus Bot. cae 16.
Evias, TS. 1977. problems in North America. An overview.
In: G.T. Prance and TS. Elias, acs, Pecado is forever. New York Botanical Garden,
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FERNALD, M.L. 1 i . Gray's manual of botany, 8th ed. Reprinted 1987. Dioscorides Press,
Portland, OR.

FLAcK, S. and E. Furtow. 1996. America’s least wanted. Nat. Conservancy Mag. 46(6):17—

aon ].W. 1983. Native distribution of the Sa/vinia auriculata complex and keys to spe-
cies identification. Aquat. Bot. 17:71—

GLEASON, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United
States and adjacent Canada, 3 vols. New York Botanical Garden, Bronx

and A. Cronquist. 1963. Manual of the vascular plants of northeastern United

States and adjacent Canada. Van Nostrand Reinhold Company, New Y
and 1991. Manual of the vascular ara e oe enue United
States and adjacent Canada, 2nd ed. New York Botanical Garden, Bre
GRAHAM, S.A 1986. gee ee In: Great Plains Flora Association. F me of the Great Plains.
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Harvey, W.D. 1998. eee or oe harmful fish, shellfish, and aquatic plants. In:

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Haren, $.L., K.N. GANDHI, and L.E. Bkown. 1990. Checklist of the vascular plants of Texas
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JACONO, C. 1998. Have you seen this plant? teas Parks and Wildlife information sheet.
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JONES, ore am and P.M. MonrGomery. 1997. Vascular plants of Texas: A com-
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Mirsky, S. 1999, — fiend: The Old World climbing fern speeds its assault on Florida.
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O'KENNON, R.J., G. Dicss, Jr., and B.L. Lipscoms. 1998a. Lactuca saligona L. (Asteraceae)
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1284 Sipa 18(4)
BOOK REVIEWS

RicHARD P. WUNDERLIN. 1998. Guide to the Vascular Plants of Florida.
(ISBN 0-8130-1556-1, hbk). University Press of Florida, 15 North-
west 15th Street, Gainesville, FL 32611, U.S.A. $35.00. 806 pp.

This is an identification guide to the marvelous vascular flora of Florida, a state whose
pant diversity is surpassed in the U.S. only by California and Texas. Of Florida’s 227 fami-
lies, 1306 genera, and — 4 species, four genera and pecies are endemic to the state.
ee 44% of the species are exotics. Genera and Res oo meee) under each
family, but the families are in poy cacaceec) se istrations are given
except fora county map of Flori lrawing FTillandsia eae: ‘fuzzywuzzy
airplant”) on the citle page. The indexes are three: common names, scientific names, and
sai names; a poland index is on cover 3 and the facing page, a helpful arrangement. The

major floristic works; no generic revisions are listed. A

n

7 literature citations are largely to
eae defines about 600 terms. Data for each species are scientific name, common name,
habitat, frequency, brief notes on in-state range, powering. one and sometimes a Hones
synonymy. (An “Atlas of Florida Vascular Plants” is availabl sf.edu/
“apc atlas.html.) The keys to groups and families occupy oy pages. of tested
them with 17 species; 13 worked well enough, but four were problematical: Erigenia bulbosa
ai Galinm aparine (sepals obsolete, but Apiaceae and Rubiaceae key out under ae and
corolla present”); acaulescent violets (Violaceae ie out under “leaves alternate”); and Najas,
1 leaves (stopped at “leaves with parallel veins”). (This is not the only
flora with this problem.) The book is well sere and pies Up co now, visitors to Florida
have ha i to take along three guides to the flora (Long & Lakela 1971; Wunderlin 1982;
985), but now one will suffice. Ac $35. ne he book is a bareaia. Kudos to Dr.
John W. Thieret.

with its single-veinec

Clewell
ve and the University Press of Florida.

Stipa 18(4): 1284. 1999

BOUTELOUA RIGIDISETA (POACEAE)
NEW TO LOUISIANA

CHARLES M. ALLEN, CHRIS S. REID, and CHRIS H. DOFFITT

Department of Biology
The University of Loutsiana at Monroe
Monroe, LA 71209, U.S.A
biallen@alpha.nlu.edu

A recent grass collection from Fort Polk, in west central Louisiana, is ap-
parently the first record for Bouteloua rigidiseta (Steud.) Hitchc. for the state.
It is not in Allen (1993) nor Thomas and Allen (1993). In Gould (1975),
the distribution is Texas, Oklahoma, and Mexico, while Smith (1994) in-
cludes this species in the Arkansas flora. The other two species of Bowtelona
in Louisiana are rare, with B. hirsuta Lag. recorded only from Calcasieu Parish,
and B. curtipendula (Michx.) Torr. from Caldwell, East Baton Rouge, Natchitoches,
and Rapides parishes. The Caldwell and East Baton Rouge collections are
recent, but the other three are more than 50 years old.

Voucher specimens: LOUISIANA. Vernon Parish: roadside at of Entrance Road
and Avenue A, ca 1 mi E of La 184 on North Fort section of Fort Polk, ca 5 mi ESE of
Leesville, Section 34 T2N R8W, 27 May 1999, Reid, Allen, and Doffitt 423 (NLU); same
locality, 25 Sep 1999, Allen 18400 (NLU).

REFERENCES
ALLEN, C.M. 1992. Grasses of Louisiana, second edition. Cajun Prairie Habitat Pres. Soc.,
Eunice, LA
GouLp, FW. 1975. The grasses of Texas. Texas A. & M. Univ. Press, College Station, TX.
SmiTH, E.B. 1994. Keys to the flora of Arkansas. Univ. Of Arkansas Press, Fayetteville,
AR.

THoMAS, R.D. and C.M. ALLEN. 1993. Atlas of the vascular flora of Louisiana, Vol.
and ferns allies, conifers, and monocotyledons. Louisiana Department of car oA
Fisheries, Baton Rouge,

Sipa 18(4): 1285. 1999

1286 Sipa 18(4)
BOOK REVIEWS

SUSANNE M. ScHutrz, AMY E. DUNHAM, Karen V. Root, SHERYL L. Soucy,
STEVEN D. Carrot, and Lev R. GinzBurG. 1999. Conservation Bi-
ology with RAMAS Ecolab. (ISBN 0-87893-768-4, pbk.) Sinaur
Associates, Inc., PO. Box 407, Sunderland, MA 01375-0407 (413-

49-4300; 413-549-1118 fax; publish@sinauer.com). $27.95. 216 pp.
Illustrated, CD-ROM disk.

Conservation Biology with RAMAS Ecolab is an excellent resource for beginning students of

applied ecology or conservation biology. It has been designed to complement most major

Conservation Biology texts, as highlighted in a chart that cross-references book chapters

with relevant lab activities. The dielass and ata ae age alias are well written and

| cechniques

provide a collection of exercises that 1
that are the foundation for these fields. Students begin with the basics of sampling tech-
niques, the use of biological indices, and describing population dynamics. In later chap-
rey explore techniques for facilitating biological management through sensitivity analyses,

ters ¢
determining Maximum Sustainable Yields, reserve design and more.

The RAMAS software that accompanies the manual is very easy to use and is a valuable
teaching tool that greatly enhances the learning potential from labs. In each exercise stu-
dents use computer simulations and models to explore the various topics. Questions ancl
straightforward activities guide che user through the computer program and help them
understand the significance of each of the model parameters. The manual not only pro-
vides the beginner with training in techniques and model use, but it encourages thought-
ful evi es and analysis on more difficult issues. T ie background information accompa-

ation biologists-ranging

a the Shomesnninn of ieee estimations to the controversy over conservation ver-
sus preservation, to the problems associated with reserve size and number. The data and
dilemmas used for the activities are representative of the diversity of issues and organisms
that conservation biologists work with, and will easily spark the interest of students new
to the field,

Contents of Conservation Biology with RAMAS Ecolab: Biodiversity: The diversity of life,
From Bacteria to Blue Whales: Growth without limits, Competition in Osprey, Fish, and
Barnacles: Limits to population growth, Wood Storks and Honeyeaters: Estimating popu-
lem of small populations, Giant Pandas: Risks

=

lation characteristics, Grizzly Bears: The prob
faced by endangered species, Hector’s Dolphins and the Red-Cockaded Woodpecker: Con-
serving dwindling populations, African Market Hunting and Tuna Exploitation: Main-

raining seen levels of harvesting, The African White Rhino: Too many for their own
good?, The Wild Ass and the Black-Footed Ferret: Reintroduction of endangered species,
Park aa Species Diversity: Lessons from Islands, Rescuing the Spotted Owl: Con-
serving species in multiple populations, Biodiversity’s Biggest Threat: Human population
growth, The Case of Patrick’s Marsh Wren: Making decisions to protect species, and Glos-

sary (no index).—Charlotte Bryant.

Sipa 18(4): 1286. 1999

Book REVIEWS 1287

Ros BrecMan. 1996. The Genus Matucana. Biology and Systematics
of Fascinating Peruvian Cacti. (ISBN 90-5410-638-7, hbk). A.A.
Balkema, P.O. Box 1675, Rotterdam, Netherlands (Fax: 31-104-413-
4947), published in the United States by A.A. Balkema Publishers,
Old Post Rd., Brookfield, VT 05036 (Fax: 802-276-3837). $75.00.
116 pp. text, 20 pp. of appendices and index, 80 color plates, 26 SEM
photographic plates with diagnostic drawings, 28 line illustrations.

This book is one in a series of horticultural taxonomic works published by A.A. Balkema,
of the Netherlands. The genus Matucana belongs to the Cactaceae tribe Trichocereae subtribe
Borzicactinae, and contains 19 species comprised of 27 taxa, including infraspecific variet-
ies and forms, at sodemic to che’ Western Andean Cordillera in Peru. Differences between
the taxonomic } of this work and that of Backeberg from the 1930s to the 1960s,
specifically as eeunrcted by Ritter (1981) are clearly evident throughout the text. This
work, specifically oriented toward horticultural as well as taxonomic users, is markedly
different from a standard taxonomic revision, owing to the recognition of four informal

“species groups” recognized on the basis —- morphology, the review of pollination and

. biology, cultivation informatio
e book is divided into 13 chapters, dee: 1) taxonomy and systematics, 2) mor-

Ree 3) geography and ecology, 4) reproduction, 5) cultivation, 6) systematics inside
Matucana-grouping of species, 7) key to all species of Matacana, 8) the haynei group, 9) the
aurantiaca group, 10) the intertexta group, 11) the paucicostata group, 12) checklist, and
finally 13) complete list of field numbers

The general chapters one through five are carefully done, with a lot of evidence obvi-
ously collected from living material in cultivation. In the systematics section, it becomes
obvious that the author follows the taxonomic species ete but unfortunately, no clear
discussions are included to justify why a certain species a particular group of synonyms.
Statements like (page 50) “To summarize, the differences between all taxa listed here as
synonyms of MM. Aaynei are too small to treat them as species,” are found in discussions afte
many of the Specs: While the author gives extensive synonymy lists, type specimens are
not listed d iene under the basionym, nor for of the synonyms. The reader is left without
knowing if types exist for all the names, if any or all have been lectotypified, etc. Listing of
the numerous womina nuda is ill-advised, and 1 ed nave left chat portion out unless they
were actually published. There is no formal generic description, nor citation of a type, al-
though ee ces to M. haynei as the oldest name in the group and the discussion afi iC

former monotypic circumscription give us the
While the author has obviously devoted years to the study of this group, I think the
systematic value of the monograph is compromised by its very non-standard format. One
is not sure if all historical s specimens have been re-studied, nor synonyms typified, and the
phylogeny portion of each species’ discussion gives us no clear picture of the salient fea-
tures that determines the author's placement of the taxon. It is abundantly clear that seed
morphology is employed to delimit the four infrageneric — but the illustrations on
page 36 lead this reader to think that the groups are difficult to recognize, especially be-
cause the seeds must be mature, and all portions of the ad must be intact for one to rec-
ognize the character
In summary, while this monograph is not a standard systematic treatment, it is an im-
portant contribution to a poorly understood, but important group of cacti. It will be a

Sipa 18(4); 1287. 1999

1288 Sipa 18(A4)

must for all concerned with the Cactaceae, with the Flora of Peru and the dynamics of the
vegetation in the Peruvian Western Cordillera. The SEM photos, color photographs, and
line drawings are helpful, as are the indices. It should certainly form part of any botanical
or horticultural library.—John_J. Pipoly I.

WALTER S. JUDD, CHRISTOPHER S, CAMPBELL, ELIZABETH A. KELLOGG, and PETER
E. Srevens. 1999. Plant Systematics: A Phylogenetic Approach.
(ISBN 0-87893-404-9, hbk.). Sinauer Associates, Inc., 23 Plumtree
Road, Sunderland, MA 01375-0407, USA. $67.95. 464 pp., numer-
ous figures and CD-ROM with over 650 color photographs and three
printable appendices.

Having recently watched again the movie, Mary Poppins, | am reminded of a phrase (no,
not “supercalifragilisticexpialidoceous”) that can be applied to this new text for undergraduate
courses in plant systematics—“practically perfect in every way.” This 1s the first such text
that is a product of teamwork and is the best to come along since George Lawrence's Tax-
onomy of Vascular Plants of 1951. The authors, all recognized leaders in plant systematics,
are former students and associates of Drs. Carroll Wood and Richard Howard. In fact, the
book was conceived while most of them were fellow graduate students at Harvard. They
wisely and fortuitously waited until they had professionally matured and macromolecular
techniques became widely applied before making the idea a reality. The result is a text-
book that is easily digested by students who have already mastered basic botany, portrays
the vitality and uncertainties of the current state of knowledge in the field, and introduces
both the principles of systematics and plants families of worldwide importance. Because
the authors are strong eee of the use of cladistic methods in both research and teach-
ing, phylogenetic concepts are integrated from the introduction to the last family treat-
ment. The goal is not the indoctrination of the conviction that cladistic approaches are
better than others. Rather, they hope to make students conversant with the current re-
searc h mene and to aedemand the rationale used to justify the taxa presen

] the sequence of topics follows a logical progression. After a brief eh is of
ne systematics in Chapter One, the second chapter provides a thorough primer of cla-
distic methods and a nen ae of cladistic, phenetic, and evolutionary schools. Here,
the distinction between grouping, naming, and ranking taxa is lucidly explained. The his-
torical meee in We jyapter Three is arranged topically and then chronologically, 1.¢.
the aa ous of aa uaeid relationships vs. development of the formation of fichier
taxa. The next t upters survey the data synthesized by systematists. Categories of mor-
phological, ena and ae structures are introduced and important terms are

fined. However, jargon is minimal (e.g., leaf shapes are reduced to four with auxiliary
modifiers such as “narrowly”), many terms are introduced only in ase diagrams, and
others are introduced and defined only in the family treatments (e.g., unique cucurbitoid
teeth, which are diagnostic for Cucurbitales). Pollination, dispersal, ie systems, and
chromosomes are discussed with specific examples from the literature to illustrate ie coe
data are used. PEON SEONS, especially those not trained in molecular systematics, will appre-

iate th of genome structure, laboratory methods, data types, data analysis,
and eee oa: as species trees versus gene trees. Chapter Six focuses on theoretic as-

Sipa 18(4): 1288. 1999

Book REVIEWS 1289

pects of species—the evolution of biodiversity, speciation, and species concepts. Case stud-
ies present the problems of applying species concepts to real plant groups. Constituting
over half the book, the family treatments are given in the Chapters Seven (nonflowering
ates and Eight iene The text ends with two appendices covering bo-
a as nomenclature and the preparation and identification of specimens.

° book i is designed to Be seudent eiendly,* Giic early inioaucron oo ineepration Sk
Pee will
conferences. The simplification of terminology will allow them to c ntrate on conce
instead of memorization. The family treatments include not only concise, parallel descrip-
tions ieenceal diagnostic features highlighted in Italics with synapomorphies in bold) but

geographically and economically important genera and unsolved research

eee Learning ay characteristics | is aoe oy ene use of por formulas and full-

page ae drawings f the Southeastern
United States. Only fe mailics, orders, aad higher taxa are ate recognized that are strongly

supported by structural and molecular analyses as monophyletic. Paraphyletic eA or
those ene sufficient study and evidence of monophyly are referred to in quotes, e.g.,

“Paleoherbs” and “Flacourtiaceae.” Students need only learn the characters that unite for-
merl closely related family pairs, e.g., Apocynaceae vs. Asclepiadaceae and Brassicaceae
VS. Cappatacese, The major families Ensen se au treatment include both tropical and
e extensive color photographs
on che CD- ROM en the serious student opportunities to reinforce an understanding
of the family characteristics.

“Practically perfect” of course implies some imperfection. There are some points the authors
may want to consider including in subsequent editions. I was surprised to find no mention
of punctuated equilibrium and related theoretical problems of higher taxa. Also, the text
would have wider appeal in the Desert Southwest (and arid South America) if conspicuous
families such as ere Prankentaceae, and Tamaricaceae were mentioned and briefly
characterized in the discussions of treated families to which they are related. Among the
photos on CD-ROM, I or like to see even saan taxonomic representation and close-
up dissections to enhance this already excellent resource.

milies are arranged by the ordinal classification propose by the Angiosperm Phylog-
eny an (APG). Thus, some taxa are supported, though strongly, only by molecular data
bur cannot be easily characterized. For example, tee sensu APG includes Malpighiaceae,
Clusiaceae, nine ag Paphos el anal Violaceae, Passifloraceae, and
Salicaceae, but not Polygal ly dry stigmas, a fibrous exotegmen, and trilacunar nodes
possibly unite an eee morphologically Some instructors may find this disconcerting.
However, one should keep in mind that the suprafamilial taxa presented in the book are
only those strongly supported such that this classification is likely to change minimally in
the future. Although higher taxa in traditional classifications (e.g., those of Cronquist and

e, both eae as appendixes on the CD-ROM) can be characterized morphologi-
saline current evidence shows many of the taxa in those classifications to be incorrect. Why
teach outdated systems to students, who can surely accommodate uncertainties in the sys-

m?

The overall quality of Plant Systematics 1 is outstanding. Instructors should eagerly adopt
this text (and keep it on their reference she think that it will lead students to appreci-
ate plant systematics as a field and prepare a for a career in oo better than any
other available for a one-semester, undergraduate course.—Roger W. Sanders, Botanical Re-
search Institute of Texas, 509 Pecan St., Fort Worth, TX 76102-4060, U.S.A

Stipa 18(4): 1289. 1999

1290 Sipa 18(4)

Georce Yarskievycu. 1999, Steyermark’s Flora of Missouri-Volume I.
Revised Ed. (ISBN: 1-887247-19-X, hbk). Missouri Department of
Conservation, P.O. Box 180, Jefferson City, MO 65201-0180; in co-
operation with The Missouri Botanical Garden Press, P.O. Box 299.
St. Louis, MO 63166. $38.00. xii + 991 pp., 194 plates of black and
white line drawings; 20 figures, including 27 individual black and
white photographs; 798 distribution maps.

re long awaited and much anticipated revision of volume one of Julian eos s 1963

T
classic Flora of Missouri (F lora) was finally published during the first quarter of 1999. Given

the outstanding quality of the Yatskievych’s treatment of the monocots in volume one, it
was well worth the wait! The book is so well written, organized, researched, illustrated,
arly done, that one must struggle to find faule. In the dedication to

thorough, and scho
Julian Sceyermark, the auchor humbly asks if Steyermark had still been present with us,
would he have approved the book’s contents? Not only do [ believe that Steyermark would
have answered with a very enthusiastic and resounding, ‘Yes!’, but Lam confident that he

would agree that this will surely be recorded as one of the greatest treatments of monocots
ever written!

Volume one of the F/ora consists of two parts: 1) and introduction, and 2) the Flora it-
self. The first part is so well written and enjoyable to read, that it delivers like a 91-page
natural history novel. Printed en this part would probably be a number one best
seller! The introduction includes: 1) a history of es bot iny in Missouri; an analysis

of changes in the state’s native aad introduced plant taxa since Steyermark’s original treat-

ment; 3) a description of the different natural ve isions and plant communities within the
state; and 4) a discussion on how the evolutionary history, geography, climate, geology,
landscape features, and anthropogenic atfects of man in Missouri have influenced the dis-
tribution and ecology of the state's flora.

The second part, a 900 page Flora, is nothing short of a masterpiece that will undoubt-
edly set a high standard of excellence for similar treatments in che fucure! Although space

would not allow a listing of all the “highligt its of this section, a few are worth mentioning.
The first and most obvious is the thoroughness, accuracy, and comprehensiveness of the
treatments of the families, genera, and s Species disc rate It should be obvious to any reader

that several sources of information were used i { the written text. These included:

1) an exhaustive review of the published and cue literature (an incredible 694
references cited!) involving the taxonomy, systematics, ecology, distribution, and descrip-
tion of the taxa discussed; 2) an examination of thousands of museum specimens; 3) com-
munication with numerous professional and amateur botanists who actively collect in the

correspondence and cooperation with leading experts and authorities of the groups

o

state
oe and 5) hundreds of hours of field investigations that were necessary to clarify the
distribution of numerous species.

I found the keys easy to use and follow, especially those of the larger or more difficult
groups (e.g., Carex, Cyperis, Eleocharis, Eragrostis, Panicum, Poaceae, and Preridophytes).
One plus for the keys is that the author painstakingly covered numerous exceptions found
in many a by lengthening the keys to cover such variations often typical for different
monocot ta

other cpoises highlight is che exquisite detail dis} oli ayed in the numerous illustra-

on all of which were newly produced for this portion of The Flora. The line drawings of

Sipa 18(4): 1290. 1999

Book REVIEWS 1291

each species depicted are some of the most accurate and detailed illustrations I have ever
observed. Although the overall quality of the illustrations of the nine different artists used
are excellent, I would do this review a great injustice if I did not separately mention the
outstanding quality of the drawings provided by Paul Nel son (Acoraceae, Araceae, Carex)
and Phyllis Bick [Agavaceae, Alismataceae, Cupressaceae, Cyperaceae (in part), ferns & fern
allies, Marantaceae, Naj le Orchidaceae, and Poaceae (in part)}. Nelson’s illustrations
are so accurate and complete that any reader will easily fall victim to modification of a
t ial, ‘bet you can’t view just one!’. His line drawings are so stunning in their
detail and accuracy that he is one artist who would actually become very successful if he
should ever decide ‘to quit his day job!’. Likewise, Bick’s drawings provide exceptionally
accurate depictions of groups many artists have a challenging time correctly illustrating
such as difficult species within the Cyperaceae and Poaceae. It is always refreshing to fin
a recent flora that has been newly illustrated, especially when the dr rawings are as well done
as with this work.

Another high point of the book is that a separate description, county distribution map,
and habitat affinities are provided for well over 99% of the taxa discussed. The only excep-
tions are for those few species that were discovered just as the book was being finalized
before going to the printer.

No review of this book’s high

ights would be complete without a comment on how in-
expensive it is, especially for a well bound hard copy with 991 pages. At $38, the book is
a steal and easily affordable, even for those on a tight budget

Although negative points about the book are hard to find, a few should be discussed.
The most obvious short coming is that it is difficule to find individual species of Carex
without knowing ahead of time what Section they are in, or similarly locating genera of
grasses without prior knowledge of their tribal affiliation. While this will not be a hin-
drance to experts working with Carex and the Poaceae, it will be problematic for anyone
who lacks the necessary expertise for such groups. Although there is an excellent index
following the literature cited section, it is very cumbersome to move back and forth through
such a heavy book. A simple solution would have been to provide an index to Carex s
and grass genera at the beginning of each respective treatment.

jon

dECIeS

—

The author's designations for abundance for some species are hard to comprehend. It is
difficule to understand how species that are known from ae one or two records in the
state (e.g., Carex atherodes, Carex laxiflora, Carex reniformis, Carex sartwellii, Carex straminea.
and Carex ene) can be given the same ‘uncommon’ designation as species known

milti

unties (€.g., Carex buxbaumi, Carex molestiforms, Carex iis ci and in some
asp

cases, represented by tens of thousands of plants in specific Rican (e.g., Car
Although overall the illustrations for the Poaceae are extr mely accurate, the nee
of only portions of some species prevents a full appreciation of the general habit for certain

taxa (e.g., Poa wolfit, Oryzopsis racemosa, Elymus riparius, Miscanthus spp., and Erianthus spp.).
One noteworthy error is that the last sentence in the discussion of Bomteloua hirsuta on
page 678 applies to B. gracilis as the latter is the rarer of the two in the state that is cur-
rently only known from one extant site in Atchison County. Another is associated with the
key to the Section Griseae and discovered by the author himself- the second parts of cou-
plet three: “lowermost pistillate scales with the bodies much shorter than the associated
perigynia (vs. as long as or longer than the associated perigynia)” should be moved to be-
come the respective second parts of couplet five because C. oligocarpa and C. planispicata
have the bodies of the lowermost pistillate scales shorter than the associated perigynia.
As currently written in the key, C. ofigocarpa and C. planispicata would incorrectly key out

SipA 18(4): 1291. 1999

1292 Stipa 18(4)
to C. llaccosperma or C. glaucodea, Additionally, it would have been helpful if the artist would
tillate scale for C. grisea and C. amphibola (rather than one

have illustrated a lowermost pis
245 of these two

situated farther up the pistillate spike) so that the illustrations on pages
species would correspond with the key on page 247 and the text on pages 247 and 252.
The following items are examples of minor errata ytal introduced taxa on page
83 should read 820 rather than 20, 2) the achene of Carex ok/ahomensis illustrated on page
355 should show two stigmas rather than three, 4) the illustrations tor Bothriochloa nha
and B. bladhii on page 593 are reversed, 4) the word ‘fertile’ in the last couplet of the key
to i eaiioe on page 140 should be replaced with ‘sterile’, 3 five or Six vee are listed
f s description of Isotria but only four leaves are depicted in the illustration of

lsotria ee. 6) Section Intermediae should be listed in the legend of Plate 41 for
twellii, and not Hymenochlaenae, and 7) Section Laxiflorae should be listed in the

in the gen

Carex sa
legend of Plate 41 for Carex blanda, and not Hymenochlaena

The negative points listed above are so minor and insignificant compared to the overall
excellence of this book, that they are nearly not worth mentioning. Given the quality and
price of this masterpiece, Steyermark's The Flora of Missouri- Volume 1, should be on the shelf
of every botanist, naturalist, and plant enthusiast in the Midwest! Additionally, it will be
a welcomed reference that will undoubtedly be in high demand throughout botanical circles
in North America and abroad. oe with Diggs et al. 1999 excellent I//ustrated Flora of
North Central Texas (see Marshall Johnston's complimentary review in the September 1999
issue of Sida), Steyermark’s The Flora if MG out Volume 1, is definitely a “must buy” botani-

cal reference! —Paul M. McKenzie

Sipa 18(4): 1292. 1999

Books RECEIVED 1293

BOOKS RECEIVED

JOHNSON, FReDeERIC S. 1995. Wild Trees of Idaho. (ISBN 0-89301-145-2,
obk.). University of Idaho Press, Moscow, ID 83844-1107, U.S.A. $14.95.
xlv + 212 pp.

MicHaAeL Kunns. 1998. A Guide to the Trees of Utah and the Inter-
mountain West. (ISBN 0-87421-244-8, pbk.). Utah State Univer-
sity Press, Logan, UT 84322-7800, U.S.A. $14.95. x + 342 pp.

Here are two more tree books for your library. The Idaho volume (8" = 10") treats 88

species plus 19 shrub species not known to reach tree size in Idaho. It has 33 pages of i

troductory text, including 10 pages of description of the vegetation of the state and a ae

of Idaho vegetation history” (Triassic to present), useful additions. The “Picture keys to
tree leaves” leads to the usual groups of species (e.g., broadleaf trees with alternate, simple
leaves), each with a list of common names. (There are no generic keys.) Thus, if che plant
in hand has simple, alternate leaves, you are lead to 19 common names. Then you turn to
the main part of the book and begin checking these 19 until you Bnd a match for your
specimen. Each species is illustrated with black-and-white pee ae of leafy twigs or
with drawings from elsewhere. Additionally, 80 color photos—some as small as a square
inch—are grouped onto eight full-page plates. Range is ae on maps of northwestern

U.S. The book has a glossary and indexes to common names and to scientific names.

Overlapping the Idaho book in the southern third of that state, che Utah volume (4.5"

x 8") describes all native and most introduced species, 219 in all. The descriptions of mor-

phology are well ae notes on habitat, uses, and cultivation are given. The black-and-

white illustrations are from various sources. Eight plates of color photos show 12 species,
half of them introduced. The introductory material considers nativity, common vs. scientific
names, tree parts, and tree arenes A 26-page “selection guide” summarizes data that
should be useful for people—both in Utah and in surrounding states—looking for a tree

species to plant in their front ca The key to trees works well. The book closes with a

glossary and an index to scientific and common names

Sipa 18(4): 1293. 1999

SIDA, CONTRIBUTIONS TO BOTANY AND
REVIEWERS FOR VOLUME 18, 1998-1999

The following individuals kindly supported Sipa through their time and
expertise in reviewing manuscripts published in volume 18, the largest published
to date. Your support is deeply appreciated.

Subscriptions. —SiDa is distributed in over 90 countries. There are 869
subscriptions, 380 domestic and 489 foreign. Brazil has the most subscrib-
ers outside of the U.S.A. Subscriptions for 2000: $27 for individuals, $50
for domestic institutions, and $60 for foreign institutions.

We thank all authors, reviewers, subscribers, and readers for your con-
tinued interest and support.—Barney Lipscomb (BRIT), Editor, John W. Thieret
(NKU), Associate Editor; Félix Llamas (LEB), Contributing Spanish Editor.

Adams, Robert P. Eckenwalder, James E. Lewis, Deborah Q. Skean _ James D.
Allred, Ke Endress, Mary Lis, Richard Smith, Alan R.
eee Wendy Farrar, Donald R. Lonard, Robert Smith, Edwin B.

n, John F. Flagg, Raymond O. Lott, Emily J. Smith, Gerald L.
be Theodore M. Ford, Bruce A. Lowrey, Timothy K. Snow, Neil
Barrie, Frec Fox HI, William E. Magrath, Lawrence K. Spellenberg, Richard
raring — A. Freeman, Craig C. Marsh, Daniel L. Stein, Bruce A
Benz, Bru Fryxell, Paul A. Mayfield, Mark Stevens, ee
Berry, Paul Gandhi, K.N. McNeill, John Strother, Joh
Bock, Jane H. Gardea, Jorge McPherson, Gordon Sutherland, ee M.
Brown, Larry E. Gordon, Doria R forgan AVI Takeuchi,
Brunsfeld, Steven J. srayum, Michael H Morris, Michael W. aylor, Chane M
Bryson, Charles T. Hall, David W. Naczi, Robert Taylor, Constance E.
Burger, William C. Harriman, Neil A. Nelson, Allan D. Terrell, Edward E
Burk, Carl J. Hartman, Ronald L. Neese, Elizabeth Thieret, John W.
Burks, Kathleen Hatch, Stephan Neill, David A. Thomas, R. Dale
Campbell, Christopher S. Hauk, Warren D. Nesom, Guy Thompson, yas a.
Canne-Hilliker, Judith M. Hempel, Alice Nicolson, Dan H. Todzia, Carc
Cantino, Philip D. Henrickson, James Perkins, Kent D. se nee F.
Carter, Richard Hodgson, Wendy Pererson, Paul M. Trock, Debra K.
Catling, Paul Holmgren, Noel H. Pilz, George E. Tucker, Gordon C.
Christensen, Knud I Ickert-Bond, Stefi anie M. Pipoly ILI, John J Urbatsch, Lowell E
Sina nan M. Johnson, Dale Poole, Jackie van der Werff, Henk
Clarke, David Johnson, ae P; Porcher, Richard D. Verhoek, Susan
Cc oe - D. ones, Gretchen Potter, Daniel Vincent, Michael A.
Clemants, Steven Jones, Stanley Pruski, John FE. Wendt, Tom
Coil, Nancy C. Kaul, Robert B. Rabeler, Richard Werth, Charles R.
Coons, Patricia Kearns, Denis M. Reed, Monique Wiersema, J.H.
Craven, Lyn aera ee H. Renner, Susanne S. Wipff, Joseph K
Crow, Garrett E. owal Rertig, Jeff offord, Eugene
Daniel, Thomas F. Kral, om. Rez ee Anthony Woodruff, Lindsa
Darwin, Steven P. Lammers, Thomas G. aa ht Eric H. Worthington, Richard
Davenport, Lawrence J. Landrum, Leslie R. Robbins, R. Laurie Wunderlin, Richard
Davidse, Gerrit Lane, Meridith A. Robertson, Kenneth R. — Yatskievych, George

awson, John Lavin, Geoffrey A Rossman, A.K. Zak, Johr
Delgado, Alfonso Le Duc, Alice Rzedowski, Jerzy Zanoni, Thomas
Diggs, a nits M. Lellinger, David Sanders, Roger W. Zarucchi, James
Dorr, Lav Lem David E Schmidt, John
: n Lersten, Nels R. Sheviak, Charles J.

Sipa 18(4): 1295. 1999

1296

Stipa 18(4)

INDEX TO VOLUME 18, 1998—1999

TITLES OF ARTICLES WITH AUT

A checklist for the vascular plants of Padre
e by LG. Necrere,
ue Maci KI T

Island National Seashor
A.D. NEtson, J.R. ie TZE,
WILBURN, and A. Day

A new oe
Ophioglossaceae) for North America by
James C, 7 ecH, ParriciA R. MANNING, and

Adder’s-tongue

ale

WARREN HERB WAGNER JR. -

A new "nets lum (Orchidaceae) species from
the Crater Mountain area in Papua New
Guinea by N.H.S. Howcrorr 1019

combination in Croizatia
(Euphorbiaceae) by L.J. Dorr 831

A new combination in Mexican Mandevilla
(Apocynaceae) by Justin WiLLiAMs 237

A new endemic Species of Mentzelra sect.
Bartonia (I 1 New Mexico by
THOMAS K. TODSEN 819

A new hybrid of Spiraea (Rosaceae) from
Oregon by WiLLIAM J. Hess and Nick A
STOYNOFE 827

A new ie of men aun aee by

» 69

Mark H. Mayt

A new

new species x film i
(Hymenoy siniueane Preridophyta) from
cide India sa C. ABbuUL HAMEED and P.V.

9

4 IAN 51

A new species of Mandevilla (Apocynaceae)
from Jalisco, Mexico by JusTIN WILLIAMS
231
A new ba eae of Phaseolus (Fabaceae) from
st-Central Mexico AYMUNDO
a es DELGADILLO ALFONSO
DELGADO-SALINAS 63

and

> ] \]

“HORS

Pipoty I and JON M. RIcKETSON 433
econd speci
a rede) en Mexico by Guy L.

s of Oritrophinm (Asteraceae:
NESoM
523
Additions to the genus Ardrsia subgenus
sol he einsinace ae) by JOHN J.
Pipoty HI and JON RIcKETSON 1145
Pee (Scropl ieee in the Ozark

=
=

Highlands by Joun
Agave - ophila Gatcia- el & E. Martinez
_ nov. (Agavaceae) by ABISAI GARCIA-
Nee y EsrEBAN MARTINEZ SALAS 627
An annotated preliminary checklist of the
dicotyledonous lianas and vines from the
Las Cruces aha SrAtIOn, Costa Rica
by ALEXANDER KRIN¢

wil

aj

/ Avdisia niainbiensts

yrsinaceae), anew species
of Ardisia subgenus Ardisia from the Chocé

Floristic Province of Colombia by Joun

J. Preory HE and ALVARO COGOLLO P. 389
Arkansas Carex (Cyperaceae): A briefly an-
notated list by Prine E. Hyarr 535

A rnoglossum ad thu / (Asteraceae ). New species

from northern Florida by Loran C. ANDER-
SON 377

ay

fenciron, anew generic name (Fabaceae,
¢ Desseeaq carl - 7 ry Pare
I
by Josepn H. Kirkpripe, JR. 815
Botanical results from the 1995 Bismarck-

Detarieae

ey

Ramu Expedition in Papua New Guinea
by W. Takeucul 752

Bouteloua rigidiseta (Poaceae) new to Louisi-
ana by Cuarces M. ALLEN, Curis S. Reip,
and CHrIs an _ 1285

C. aryOpsis |

A new species of P, ielic e
James HENRICKSON anc
Drvenper LOO]

A new

(R : C

jan

HOMAS R. VAN

variety of Physaria newberryi
New Mexico by Roper

C. SIVINSKI 673

A new variety of Prras greegis (Pinaceae) in

Mexico by Jerrrey K. DONAHUE and JAVIER
Lopez Upton 1083

A revision of the genus Ardssia subgenus

Graphardisia (Myrsinaceac) by JOUN J.

2906.

Stipa 18(4); 12 1999

Lepto. ile sensu lato
ace iceae, C Lia ae by Nein SNow 27

Clusia niambiensis (Clusiaceae), a new species
fron shoc6é Floristic Province of Co-
lombia and Ecuador by Joun J. Pipory IH,
ALVARO COGOLLO P., and Marra Soria
GONZALEZ 395

Comments on the distribution of Sedwm
eg hellum Se eneer in Texas by H.L.

,W.C. Hoimes, and

=)

ie eee 622

INDEX

Cyperus fuscus (Cy peraceae), new to Missouri
Nevada, with comments on its occur-
rence in North America by PauL M.
McKenzie, Brapd Jacoss, Cuaris T.
BRYSON, CORDON C. Tucker, and Ricu-
ARD CAR
nee “of the Indo-Malesian genus
Hymenandra (Myrsinaceae) in the
eotropics, and its boreotropical impli-
cations by JOHN J. Pipoty III and Jon M.
RICKETSON 701
Documented chromosome numbers

ani
ection
ve oar Additions, variations,
by P.E. Rorurock and A.A.
REZNICEK 587
Documented chromosome numbers 1999:1.
Chromosome numbers and pollen diam-
eter variation in Thelesperma (Asteraceae)
by Ler F. Greer and A. MicHaci Powe.
911
Dos nuevas adiciones a la sear en
Mexicana by ADOLFO Esprjo SERNA,
Rosa L6prz-FERRARI, JAVIER GARCIA i
ROLANDO JIMENEZ MAcCHORRO y Luis
SANCHEZ SALDANA 41 1
Eight novelties in Abildgaardia and Bulbostylis
(Cyperaceae) from South America by RoBerr
KRAL and ss K T. STRONG 837
First repor the genus Burmeistera
(Cam ae from Honduras by Trtomas
G. LAME RS and ES M. Maas 363

Chromosome numbers in Carex

I

1297

Glaucinm cornicilatum (Papaveraceae) in Texas

KIRKPATRICK and J.K. WILLIAMS 347

Habranthus tubispathus (Liliaceae) new to the
flora of Alabama by ALvIN R. DiaMonp,
JR., CHARLES P. CHAPMAN, a BRUMMETT

.
3

oy ZOE

Hoventa dulcis (Rhamnaceae) naturalized in
central Texas by DouGLas H. GOLDMAN 350

In Memoriam—Amelia Anderson Pea
(19??-1998) pyd RuTH ANDERSSON May 62

In Memoriam— z Dietrich Lubrecht
(1908-1997), botanical bookseller and

1 by Witttam R. Burk 371

Ipomoea asarifolia (Convolvulaceae), another
potential exotic pest in the United States
by KATHLEEN CRADDOCK Burks and DANIEL
F. Austin 1267

— saligna

A tél aceae), a lettuce new for
is by Roperr J. O’KENNON, GEORGE
_ De. JR. sand Barney L. Lirscoms 615

Laf SARA COMMAS

Asteraceae) new for Texas
and notes on invasive exotics by ROBERT
O’KENNON, THEODORE M. RKLEY,
Grorce M. “e Gs, JR., and Barney L.
LipscoMB 1277
Luziola pernviana (Poaceae: Oryzeae) previ-
ously unreported from Texas and a key to
Texas species by STEPHAN L. Harcn, ei
J BOGAN N, and Jue s A. THomas 61
ee weedy
in Texas by MoniQur DuBruLeé REED and
Fi aa Ket TCHERSID 3

—~

Flavonoids in tropl 1 the related
genus ae cease. Fabaceae):
Distribution and phylogenetic significance
by JUAN PaBLo Petorro and Maria A. DeL
PERO Martinez 2

neal en ple azas SY) jeri publicos

li México
by ALEJANDRA ROCHA ESTRADA, TERESA E.
TORRES CEPEDA, Ma. DEI

FONZALEZ DE LA ROSA. SALOMO

CONSUELO
YJ. MARTINEZ

Lozano Y Marco A. ALVARADO VAZQUEZ
DES

Sp ala simplicicaulis (Asteraceae:
rnaphalieae) in four southeastern states
and new a North America by Guy L.

Nesom 1259

Sipa 18(4): 1297. 1999

1me- a (Poaceae:
species from Nuevo
Leén, Mexico by Paut M. Pererson and
Jesus VALDES-REYNA 68

Me ae anew

Myrsine luae Ce eis anew ae from
razil by JON M. RICKETSON and JOHN J.
Pipoty HI _ 7
New and note worthy angiosperm records for
Iowa by Craic C. FREEMAN 605
New combinations in Setaria (Poaceae:
Paniceae) by Witiiam E. Fox, III and
STEPHAN L. Hatcu 1037
New combinations in the Melastomataceae
from Hispaniola by HENRI ALAIN Liogier
1025
New flowering plants from southern New

1298

Ireland, Papua New Guinea a. W. TAKEUCHI
and JOHN J. Piroty HI 1

New names and eae in neotropical
oe ears i) JOHN J. Pipoty IL and Jon
M. Rick N 503

New plant a for Dominica, Lesser
Antilles by SrevEN R. Hitt and ARLING-
TON JAMES 297

New plants from Crater Mt., Papua New
Guinea, and an annotated checklist of the
species by W. TAKEUCHI 941

New plants from Papuasia. Novelties from
the Lakekamu and Bismarck-Ramu expe-
ditions by W. TAKeEucHI 107

New species and nomenclatura
(Clusiaceae) from Andean Colombia and
Venezuela ere J. Preory Hand Alvaro
CoGoL_Lo P. ¢

New species of : ie (Clusiaceae) from the
Cordillera Occidental of Colombia JOHN
J. Pipory Ill and ALVARO CoGoLLo P. 1175

Nomenclatural Nuphar
(Nymphaeaceae) by DONALD J. PADGETT
823

Notes on sectional delimitations in Erigeron
(Asteraceae: Astereae) by Guy L. Nesom

nd Richard D. Noyes 1161
Notes on the flora of Texas with additions

me
—

notes in Clusia

novelties

and other significant records by Larry E.
Brown and Sruart J. Marcus 315

Noteworthy records of Mississippi vascular
plants by Bruce A. Sorrie and Steve W
LEONARD 891

Noteworthy vascular plants from Arkansas
by Eric SUNDELL, R. DALE THoMas, Cari
AMASON, Roserr L. Sruckey, and JOHN
LOGAN 879

Novelties in the Myrsinaceae from the Ven-
ezuelan Guayana by JOHN J. Pipoty HI and
JON RICKETSON 1167

Orthodon vs Mosla (Lamiaceae) by New A.
HarrIMAN 93

Packera subnuda con nov. a corrected name

Senecioneae)

A K. Trock and THEOpoRE M.
KLEY a
Paaderia foetide (Rubiaceae), new to the flora
of North Carolina by Perr DIAMOND 1273
1298.

Stipa 18(4):; 12 1999

Stipa 18(4)

Pennisetum advena sp. nov. (Poaceae: Paniceae):
A common ornamental grass throughout
the southern United States by Josepy K.
Wipre and J.P. VeLpKampP 103

Puan metcalfei (Scrophulariaceae), a valid

ecies by THOMAS K. TODSEN

Plena cor a (Plantaginaceae) new to Texas
by Rosert J. O'KENNON, GeorGe M. DIGGs,
yr., and RONALD K. HoGGarb 356

Pollen morphology of the genus sala il
(Cucurbitaceae) by CONCEPCION RODRIGUEZ
J. and RODOLEO PALACcIOs-CHAVEZ ae

Agalinis harperi

Ww

Priority of the name
(Scrophulariaceae) over the names Agalinis
delicatula and Agalinis pinetorum by JOHN
F. Hays 369

Preris vittata cineridles eae), a new fern for Texas

2M. Dice

Jr. 359
Review of early nomenclature in Evthamia
(Asteraceae: Astereae) by Guy L. Nesom
)O9
Ruellia jimulcensis (Acanthaceae), a new spe-
cies from the Chihuahuan Desert area,
Mexico by José A. VILLARREAL Q. 223
Salvinia molesta (Salviniaceae), new to Texas
ind Louisiana by CoLeTre C. JACONO 931
ee rosea (Sarraceniaceae), a new spe-
cies of pitcher plant from the southeast-
ern United States by Rosert F.C. Nacz1,
ric M. Soper, FREDERICK W. Case, JR.,
ina Roperta B. Case 1183
Seasonal changes in concentration and dis-
tribution of heavy metals in creosotebush,
Larrea tridentata (Zy gophyllaceae), tissues
in the E] Paso, TX/Ciudad Juarez, Mexico
area by WILLIAM P. Mackay, RICHARD MENA,
NICHOLAS E. PINGITORE JR., KEITH REDETZKE,
C. Epbwarb FREEMAN, HAROLD NEWMAN,
ees isan A, and Herc TOR ~~ 287
-

Sort

Che AD FESYCO

(eins harniacess) with erect growth hab-
ics by W. JoHN Hayben and OLGa
TROYANSKAYA 419
Seven new nomenclatural combinations and
in Packera (Aste
Senecioneae) by Desra K. TRock and
THEODORE M. BARKLEY 38

a new Name raceae:

INDEX

triking sexual dimorphism in Lindera
subcoriacea (Lauraceae) by Loran C. ANDER-
son LO
Studies in New World Amaranthus
Salam ge by JAMES HENRICKSON 783
Synops World Commicarpus
cee by Net. A. HARRIMAN 679
Taxonomic clarification of Atriplex nuttallii
rapier ais and its near relatives by
ARD C. Sturz and eae C,
Sues 193
Taxonomy of the Sporobolus ean com-
plex (Poaceae: See es e) by ALAN S
WEAKLEY and Paut M. as 247
The flora of Indian shell rings
South Carolina to northern Florida by
RICHARD Sree Maritess Leyva, and
DwiGHt T. KINCAI
The genus ae eee in Ec-
uador a . Pipoty UI 1
The genus Myrszne eee ae in Venezuela
by JON M. RICKETSON and JOHN J. PIPOLY
III 1095
The oe of the American black-fruited
: ns C see s erythropoda, C. rivilaris,

from coastal

ee and C. brachyacantha to C. se
Douglasianae (Rosaceae) by J.B. PHipps ye
The segregates of Senecio, s.1., and Cacalia,
s.l., in the flora of North
of Meascicg i THEODORE M. BARKLEY 661
Cvus VIVeNS ne on ECVuUS

America north

et
drummondit Cae eae) in the southeastern

tates by RICHARD Carter, Davip
IXANDER, CHARLES T. BRYSON, and

nited

The use of animal-dispersed seeds and fruits
in forensic botany by Barney L. Lipscoms
and GeorGce M. Dicas, Jr. 335

The vascular flora of Biscayne National Park,

orida by RICHARD STALTER, JOSEF TAMORY,
PATRICK LYNCH, and Brian Lock woop 1207
Three new species of Crataegus (Rosaceae) from
ern North America: C. okennonit, C.
ee and C. phippsii by J.B. Puipps
and R.J. O’KENNON 169

Sipa 18(4): 1299. 1999

L299

GB socie (PD re \

A new species from Belize by JosepH K.
Wiprr and StaNELY p _ JONE s 241

Triosteum (C Texas by WALTER
C. Hous, Kay M. FLeminGc, RutTH and
Ron Lope d Jason R. SINGHURST 929

Triraphis sn Cae re a species
reported new to the United States by
STEPHAN L. Hatcu, WILLIAM E. Fox III,
JOHN E. Dawson III, and WitiiaM B.
GODWIN 365

Two new species of Tripogon pega from
India by A.K. Fea DEEP and C NIL iis

ave. §i ee Litta

Una
(npavaces) de Guerrero y Oaxa xaca, México
by ApisaAf GARC{fA-MENDOZA and EsTEBAN
Martinez SALAS 227

Una nueva especie de Polianthes (Agavaceae)
del estado de Oaxaca, México by ELoy
SOLANO CAMACHO and ABISAI GARCIA-
MENDoza 473

Uniqueness of the endangered Florida Sema-
phore Cactus (Opuntia corallicola) by DANIEL
F. Austin, Davip M. BINNINGER, and
one Pica I27

Variation in the Berlandiera pumila (Asteraceae)

mplex by G.L. NesoMand B.L. TURNER
493

Vascular plants new to Kansas by Cralic C.
FREEMAN and RONALD L. McGREGOR 593

Vascular plants on a gypsum outcrop in south-

exico: A listing, a new vari-
and taxonomic realignments in the
complex

Anulocaulis leiosolenus
(Nyctaginaceae), and a
Mentzelia humilis (Loasaceae) by RICHARD
PELLENBERG and THOMAS WOOTTEN 987
Vicia lutea (Fabaceae) new to Texas by AMANDA
K. New 1265

J} bear ay roe
LELASCHOTLE

new variety of

Senecioneae): A new

<—

genus and combination from Mexico by

ONNIE L. CLARK
Xanthostemon poo (Myrtaceae), a new
s from the Philippines by Peter G.

WILSON and seine L. Co 283

1300 Stipa 18(4)

AUTHOR INDEX

Alexander, David K. 1049 Harriman, Neil A. 679, se
Allen, Charles M. 1285 Hatch, Stephan L. 305, 611, 1037
Alvarado Vazquez, Marco A. 579 Hayden, W. John

Amason, Carl 879 Hays, John FE. DD
Anderson, Loran C. 1065 Henrickson, James 783, 1001
Austin, Daniel E. 37, 1267 Hess, William J. 827

B.L. Turner 493 Hill, Steven R. 297

Barkley, Theodore M. a 661, 861, 1277 ae Ronald K. 356
Binninger, David M. Holmes, Walter C. 622, 929
Branch, J.R. 622 Howcrott, N.H.S. LOL9
Brown, Larry E. 315 Hyace, Philip E. 535
Brummect, Jim, 353 Jacobs, Brad 325

Bryson, Charles T. 325, 1049 Jacono, Coletre C. 931

Burk, William R. 371 James, Arlington 297

Burks, Kachleen C aoe 1267 Jiménez Machorro, eal All|
Carter, sees 325, 1049 Jones, Stanley D. 2:

Case, Jr., Frederick W. 1183 Ketchersid, Mary ae

Cage, ae B. 1183 Kincaid, ee T. 863
Chapman, Charles P. 353 Kirkbride, Jr., Joseph H. 815
Clark, Bonnie L. eee k, Zoe 347

Co, Leonardo L. Kral, Robert 837

Cogollo P., Alvaro oo. 395, 401, 1175 Krings, Alexander 1247
Dawson ee John E. 365 Lammers, Thomas G. 363
Day, A. 1227 Lazari, Andreas 1049

Delgi oe ane as, aces 637 Leonard, Steve W. 891
Diamond, Jr., TR. 353 Leyva, Maritess 863
es ae oe Liogier, Henri Alain 1025

8, Jt. George M. 335, 356, 359, O15, Lipscomb, Barney L. 335, 615, 1277

Lockwood, Brian 1207

okie. Chris H. 1285 Logan, John 879

Donahue, ras K. 1083 Loper, Ron 931

Dorr, L.J. 831 Loper, Ruth 931

Espejo Serna, Adolfo 411 Lopez Upton, Javier 1083
Fleming, Kay M. 929 Lépez-Ferrari, Ana Rosa 411

Fox HI, William E. 365, 1037 eae Patrick 1207

Freeman, C. Edward 287 Maas, P.J.M. 363

Freeman, Craig C. 593, 605 Mackay, William P. 287

Garcia Cruz, Javier, 411 Macke, L. 1227

eee e Abisaf 227, 473, 627 M:z pdhiicoodanan: PV. 519
sardea, John 287 Manning, Patricia R. 307

eae in, William B. 365 Marcus, Stuart J. 315

Goetze, J.R. 1227 Martinez Lozano, Salomén J. 579
Goldman, Douglas H. 350 Martinez Salas, Esteban 227, 627
Gonzalez de la Rosa, Ma. del Consuelo 579 May, Ruth Andersson 629

Greer, Lee F. 911 Mayfield, Mark H. 693

Hameed, C. Abdul 519 McGregor, Ronald L. 593

Sipa 18(4): 1300. 1999

AUTHOR INDEX

McKenzie, Paul M. 325

Mena, Richard 287

Naczi, Robert F.C. 1183

Navarro, Hector 287

Negrete, I.G. 1227

Neill, es = oS

Nelson, A.D.

Nesom, Guy - re 523, 1009, 1161

Newman, Harold 287

Noyes, Richard D. 1161
)’Kennon, Robert J. 169, 356, 615, 1277
Pablo Pelotto, Juan 2

Padgett, Donald J. 82:

Palacios-Chavez, Rodolfo 479

Pero Martinez, Marfa A. Del 213

Peterson, Paul a 247, 685

Phipps, J.B. 169, 647

Pingitore Jr., ae E. 287

Pinkava, Donald J. a

, 1259

ee : = ae Jaga lis 389, 395, 401,
1136, 1167,

FOL. oo toe

1]
br A. Michael 911
Pradeep, A.K. 809
Ramirez- ect Raymundo 63
Redetzke, Keith 2
Reed, aie Dab 361
Reid, Ch
eee er
Ricketson, ae ae 503,701, 747, 1095,
1145
Rocha Estrada, Alejandra 579
Rodriguez J., Concepciédn 479

Sanchez Saldafia, Luis 411
Sanderson, Stewart C. 193
Singhurst, J.R. 622, 929

Sipa 18(4): 1301. 1999

Sivinski, Robert C., 673
Snow, Neil 271
Sofia Gonzalez, Marta 395
Solano Camacho, Eloy 473
Soper, Eric M. 1183
Sorrie, Bruce A. 891
Spellenberg, Richard 987
Stalter, Richard 863, 1207
Stanford, Jack W. 359
Stoynoff, Nick A. 827
Strong, Mark T. 837
Stuckey, Robert L. 879
Stutz, Howard C. 193
Sundell, Eric 879
Sunil, C.N. 809
Takeuchi, W. 161
Tamory, Josef 1207
Thomas, James A. 611
Thomas, R. Dale 879
Todsen, Thomas K. 621, 819
Torres Cepeda, Teresa E. 579
Trock, Debra K. 385, 861
ae be Olga, 419
ucker, Gordon C. 325
She ere 685
Van Devender, che R. 1001
Veldkamp, J.E. 1
Vall arreal Q., ak —
Wagner Jr., Warren Herb 307
Weak! me Alan S. 247
White, H.L. 622
cin T. 1227
Williams, Justin 231, 237, 347
Wilson, Peter G. 283
Wipff, Joseph K. 241,
Wootten, Thomas 987
Zech, James C. 307

1031

, 752, 941, 1071

Sipa 18(4)

1302
BoTaNIcaL NAMES
New names in bold face
Abildgaardia reflexus 993
baeothryon 839 Arabis hirsuta var. adpressipilis 594
mexicana 839 Ardisia 1145
ovata 839 alajuelae 511
papillosa 842 albipedicellata 511
Abronia ameliae 630 albipetala 515
Acalypha deamii 593 albisepala 511
Agalinis 555, 557 albovirens 468
aspera S60 apodophylla 509
delicatula 369 azaharensis 511
fasciculata 561 barclettit
gattingeri 563 SuPSP: bartlettii 461
harperi 369, 370 subsp. lilacina 466
heterophylla 563 bekomiens )9
homalantha 565 bristani 51]
pinetorum 369 chiriquiana i 1
purpurea 567 dressleri 511
skinneriana 570 dryeri 512

cenuifolia 572 duripetala 515
viridis 574 eciliata 512
ellipticifolia 512

Save
dussiana 299 eucuneata 512
gracilis 2 eurubiginosa 509
petrophila 627 feniana 512
Agrimonia incisa 891 guanacastensis 512
Alternancthera sessilis 316 guinealensis 512

Alysicarpus vaginalis 31 hatoana 515
Alyssum desertorum var. desertorum 393 heterotricha 512

Amaranthus 783 hirsutissima 510

californicus 594 hornitoana 513
crassipes var. crassipes 786 hugonensis 513
var. warnockit 788 hyalina 1158
polygonoides 797 ixcanensis 513
scleropoides 790 jaliscensis 513
tamaulipensis 800 knappit 513
xtexensis 791 lajana 515
xtucsonensis 804 latisepala 513
Antennaria parvifolia 594 leptopoda 513
Anulocaulis 987 mesoamericana 515
annulatus 992 nonteverdeana 513
eriosolenus 992 ae 513
hintoniorum 991 neomirandae 514
letosolenus 992 nevermannii 510
var. gypsogenus 992 niambiensis 390
var. howardiit 989 oaxacana 314

Sipa 18(4): 1302. 1999

BOTANICAL NAMES

opegrapha
subsp. opegrapha 443

parvidenticulata 514

parvipunctata 514

quadrata 509
malana 514

sordida 509

squamata 509

steinii 514

stevensi 51

subgenus Graphardisia 433, 436, 1147

tartariae 510

toroana 514

verapazensis 1148
subsp. cucullata 1156
mae verapazensis 1150
warneri 515
webedbauet 438

tuberculosa 892

condensata 892

simpliciflora 892
Armoracia lacustris 316, 594
Arnoglossum 665

um 378

Aster lateriflorus 595
Astragalus distortus var. distortus 606
Atriplex

cuneata 203

falcata 205

gardneri 206

nuttallii foe 203

prostrata 595

ee 209
Avicennia germinans 304
Barkleyanthus 664

riedeli
Berberis oe 595
Berlan

cde ai 497
pumila 493, 499
var. scabrella 499
Stipa 18(4): 1303. 1999

Besseya bullii 606
Bouteloua rigidiseta 1285
Brachionidium parvum 300
romus nottowayanus 595
Bublostylis carajana 844
Bulbophyllum 1019
foetidum 1019
var. oo 1019
wakoi 1019
Bulbosty "
eleocharoides 846
latifolia 848
lombardti 850
pachypoda 852
scirpoides 855
spectabilis 857
Burmannia biflora 893
Burmeistera virescens 363
Calibrachoa parviflora 596
Calopogon multiflorus 893
Cardamine
concatenata 878
debilis 317
debilis 878
_ flexuosa 879

conoidea 606
feta 590

foenea 590

hyalina 591

longti 591

merritt-fernaldii 591

molesta 5

muskingumensis 591

oronensis 591

praticola 59]

serantica 592

siccata 607

tetrastachya 592
Centraurium texense 881
Chamaecrista deeringiana 893
Chamaesyce

hirta 419

hypericifolia 419

mesembrianthemifolia 419
Chenopodium pumilio 596
Cissus obovata 304

1303

Sipa 18(4)

1304
Cladium mariscoides 893 flavovirens 13
Clidemia 1026 fonteo ; 4d
Clinopodium gracile 881 | | subsp.magnoliifol
Clusia 1175 64
aristeguictac 408 fuscus 149
bernardoi 402 gigantophyllus 30
comans 408 grandezit 139
deminuta 1175 granulosus 133
divaricata 408 guyanensis subsp. pseudoicacoreus 49
epiphytica 408 huampamiensis 30
mirandensis 408 humilis 74
niambiensis 396 incognitus 125
paisarum 1179 jensonit 142
sect. Havetiopsis 408 kayapii 84
sect. Oedematopus 408 laetus 41
tetragona 404 lepidotus 39
Coelorachis cylindrica 893 liesneri 1168
Commicarpus lineatus 117
coctoris 679 magnus 118
pretermissus 68 | subsp. asymmetricus 120
Conocarpus erecta 30] subsp. magnus 119
Convolvulus nodiflorus 302 marginatus 114
Conyza bonariensis 878 minuciflorus 127
Corsia ene var. wiakabui 164 nanayensis 109
Crataegus nestorii 39
ou. icantha 647, 658 occigranatensis 33
douglasiit 175 pastensis 24
erythropoda 647, 652 penduliflorus 153
okanaganensis 178 peruvianus 43
okennoniu 170 poeppigi 98
phippsiu 185 pseudolongifolius 101
rivularis 647, 652 resinosus 147
saligna 647, 656 schlimii 92
series Douglasianae 674 simplex 80
Series Purpureofructi 184 spicatus 61
williamsit 18] spichigert 36
Crepidomanes indicum 520 sprucei 76
Crepis setosa 878 subgenus Comomyrsine 66
Croizatia brevipetiolata 832 subgenus Conomorpha 28
Cuphea carthagenensis 882 subgenus Cybianchus 121
Cuscuta polygonorum 317 subgenus Grammadenia | 1 2
Cybianthus | subgenus Laxiflorus 58
ancthuriophyllus 89 subgenus Microconomorpha 23
cenepenis 106 subgenus Triadophora 92
comperuvianus 47 subgenus Weigeltia 97
croatit /| timanae 33
Cuatrecasasil 35 vasquezit 103
cyclopetalus 151 ree 137

verticilloides 69

Sipa 18(4): 1304. 1999

BOTANICAL NAMES

Cyperus 1049
cuspidatus 880
deormmonall 10m; 1059
eragrostis
fuscus 32
virens 1049, 1055
Dalbergia monetaria 302

2;

Dichanthelium

erectifolium 894

fusiforme 894
Didiplis diandra 882
Digitalis lanata 596
Diplacrum capitatum 299
Dirca palustris 596
Dolichopsis 213

paraguariensis 215
Dryopteris ludoviciana 894
Echinocystis poe 486
Echinopepon

cirrho aca 481

coulter! 482

gemellus 482

wrightii 482
Elaeagnus umbellata 597
Eleocharis
compressa 318
elongata 894
intermedia 605
melanocarpa 895
robbinsii 895
Elyoneurus tripsacoides 895
morya rinconensis 694
Eodea bifoliata 597
Eragrostis oe 597
aaa 11
162
ae 1161
sect. Geronpternix 1163
lepidopodus 1163
rhizomatus 1163
sect. Lamprocaules 1162
sect. Linearifolii 1162

Sipa 18(4): 1305. 1999

1305

sect. Pycnophyllum 1162
ochloa acuminata var. acuminata 883
Erychronium albidum 882
Euonymus fortunei 598
Eupatorium eens var. holzingeri 598
Euphorbia balbisii 3C
Euthamia 1009, 101 16
caroliniana 1016
galetorum 1016
graminifolia 1017
leptocephala 1017
occidentalis 1017
Fatoua villosa 883
Fimbristylis cymosa subsp. s ae 300
Floerkea proserpinacoides 6
Forestiera segregata 303
Gamochaeta simplicicaulis 1259
Gentlea
austinsmithiit 508

micranthera 508

standleyi 5

venosissima 509
Geum vernum 60
Glaucium corniculatum 347, 348
Glochidion beehlerti 945

Gordonia lasianthus 896

Habranthus tubispathus 353
Hasteola 666

—

QS

Hovenia dulcis 35(
Hybanthus concolor 886
Hydrocotyle sibthorpioides 878
Hymenandra 701

acutissima 738

callejasii 736

calycosa 7 8

pittieri 731
sordida 722
squamata 725
stenophylla 715
wilburiana 728

—

Ichnanthus nemorosus var. swartzii 300
Ipomoea asarifolia 1267

Isachne

1306

angustifolia 300

arundinacea 300
Isoetes

louisianensis 896

melanopoda 897
Jasminum pipolyi 948
Juncus gymnocarpus 897
Lactuca saligna 615
Lapsana communis 1277
Larrea tridentata 287
Leandra 1025

humilis 1026

hybophylla 1026

ees aie 1026

polychaeta 1026
Lepidium cin 879

pot

Leptochloa 2
uninervia 598
virgata 301
Lespedeza bicolor 599
Lilaeopsis chinensis 316
Lindera subcoriacea 1065
Lindernia crustacea 322
Lobelia boykinit 897
Loeflinga squarrosa 599
Ludwigia microcarpa 320

Lupinus argenteus var. argenteus 599
Luzi

peruviana a 1,612
Lycianthes asaniolia 361
Lysimachia japonica 884
Magnolia macrophylla 882
Malaxis

alvaroi 414

palustris 412

coronaria var. coronaria 599
floridunda 599

rah

gilensis 486
macrocarpa 486

Marshallia trinervia 898

Marsilea polycarpa 299

Sipa 18(4): 1306. 1999

Sipa 18(4)

Medinilla
bismarck- “ramuensis 1071
sapoi-riveren
Me ies anaes ee
Mentzelia 98
conspicua 819
humilis 994
var guadalupensis 995
jemezensis 994
multiflora 994
strictissima 994
Mitreola oe 898
Mosla 926
Mullen jaime-hintonii 686
Murdan
tied 8
se 879
lum laxum 898

Myriophy

oN

Myrsinaceae |
Myrsine L095
andina | 122
cortacea 1108
subsp. coriacea 1109
subsp. reticulata 1115
dependens 506, 1103
guianensis L136
juergenseni 506
luae 747
macrocarpa 1129
maguireana : 116

minima

no ae

nitida 113:
pellucida 1131
alee aneteoe 506
perpaucifle 12
picturata 7
resinosa 1124
schomburgkiana 1141
Nuphar 823
advena subsp. orbiculata 824
subsp. ozarkana 82
subsp. ulvacea 825
pumila subsp. sinensis 825
bsp.oguaraensis 825
sect. Astylus 824
Nymphaea amazonum 303
Ophioglossum
igelmannit 309
polyphyllum 309

BOTANICAL NAMES

Opuntia

corallicola 527, 531, 532

millspaughii 532

nashii 532

rubescens 532

spinosissima 532
Oritrophium durangense 523
Orthodon 929
Ossaea rubrinervis 1029
Packera 6

bolanderi var. harfordii 386

ueku 387
dimorphophylla var. intermedia 386

var. paysoni 386
neomexicana var. metcalfei 386
var. toumeyi 386
— var. flavula 386
var. semicordata 386
eda 635
Paederia foetida 1273
Parasenecio 665
Parnassia grandifolia 899
Paspalum dissectum 600

macrostachys 1031
orientale 1031
setaceum 1031

Penstemon metcalfei 621

Pericallis 666

Pharus oo 301

Phaseolus albescens 638

Phragmites australis 301

pudens 880
tenellus 880
Phyllocarpus 815
Physaria newberryi var. yesicola 673
Pilea hedemarkii 770
Pilocarpus racemosus 303
Pinus 1083
gregegii var. australis 1083, 1092
sca neh 356
Platant
ee vat. esl 607
psycodes

SipA 18(4): 1307. 1999

Polianthes bicolor 473
Polygala en 899
Polygon

Cespitos

cespitosum \ var. longisetum 600

umbraticola 884
recorensis 1002

y
Potamogeton epihydrus 899

Potentilla intermedia 608
runus gideonti 765
Psacalium 665
Pseudogynoxys 665
Psilotum cain 884
Psychotria
howcroftii 767
matambuai 950
osiana 162
Pteris
multifida 884
vittata 359
Pyrus calleryana 885
Randia nitida 303
Rapane
sie 506
duidae 1141

Raphanus sativus 879
Rhamnus cathartica 600
Rhynchosia minima 880
Rhynchospora

curtissi1 899

debilis 318

decurrens 898

fascicularis var. distans 900
gle larga var. pinetorum 900

har peri 90(

Roldana 664
Rorippa sylvestris 879

Rottboellia cochinchinensis 883

1307

1308

Ruellia
jimulcensis 223
eee Lssp. . pinetorum 901

Rugelia 665

Rumex
paraguayensis 322
scenophyllus 605

Saccharum coarctatum 321

Sagittaria isoetiformis 902

Sagraea 1046
abbott 1027
barahonensis 1027
cinerea 1027

lanceifolia 1028
oligantha 1028
polychaete 1028
pusilliflora 1028
setulosa 1028
woods L029
Salvinia molesta 927
Sanguinaria eartadensis 883
Sarracenia 1183
purpurea 1183

ee)
co

rosea |
. luteola 1192
a se seas eliormeds 600
Scirpus cubense 319
Scleria reticularis 902
Scutellaria racemosa 881
Sedum pulchel lum 623

pumila 883
reverchonti subsp. firmula 1042
subsp. ramiseta 1041
ebee Reverchontiae 1039
utowanaea var. eee 1043
painter 10
Shee argentea 60
sida
jamaicensis 303
rhombidfolia 601
Sinoseniceo 664
Sisyrinchium exile 88]

Stipa 18(4): 1308. 1999

Stipa 18(4)

Sorghastrum apalachicolense 902

Spartina patens 30]

Spergularia marina 606

Spiraea x hitchcockii 827

Spiranthes brevilabris var. floridana 90

Sporobolus
curtissi 253
floridanus 247, 256
pinetorum 258
silveanus 261
teretifolius 262

Stella
vallida 601
parva 317

Strophostyles 213
helvula 21

eiosperma 215

umbellata 215
Stylisma

aquatica 9OS

pickeringii 904

Stylogyne viridis 117

Tephroseris 664
Thelesperma 909
Torilis arvensis 340
Trachypogon
mayaénsis 2: 42, 244

Tradescantia crassula 879
Trautvetteria caroliniensis 884
Trichomanes crinitum 299
Trichospermum stevensti 1077

ridens

fo)

yak lahomensis 60 |
carolinianus 905
Trifolit

incarnatum 602

striatum S80
Triosteum

angustifolium 931

perfoliatum 931
Tripogon

sivarajanit 809

vellarianus 811
Triraphis mollis 365
Urochloa

)]

plantaginea 3

ramosa 321

BOTANICAL NAMES 1309
Utricularia Vici

foliosa 320

olivacea 906
Valerianella locusta 885

hirsuta 88 1
lutea 1265
Villasenoria 632
Be _

Veronica ttii 633
biloba 60 Xanthostemon fruticosus 283
hederifolia 602 Yermo 664

persica 602 Zunilia 1146

SUBJECT

Adder’s-tongue 307 cea Mexico 411
Alabama 353, 1183, 1273 Gu ero, Mexico 227
Amelia Anderson Lundell 629, 630 ypsum oye! 5 087
Animal-dispersed seeds and fruits 335 Hiwiior s 64
Arizona 802, 804 Heavy aa 287
Arkansas 535, 877 Heinz Dietrich Lubrecht 371

D

jon

Belize 241
Biscayne National Park, 1207 :
Bismarck-Ramu, Papua New Guinea 71, 1071 India 519, 809
Bog Spice Bush 1065
Boreotropical implications 701
ntral Texas 350
Chihuahuan Desert 223, 783
Choc6 Province 389, 395
Chromosome numbers 587, 909
Coahuila, Mexico 693, 783
Coastal South Carolina 861
Colombia 389, 395, 401, 1175 Las Cruces Biological Station, Costa Rica 1247
Cordillera Occidental, » Colombia 1175 Las Orquideas National Park, Colombia 1175
Costa co Las Cru Louisiana 927, 128
Crate papas sue Guinea 941, 1019 Mexico 223, 237,411
oo h 287
Dominica, Lesser aoe 297
Durango, Mex
Ecuador 1, oe
El Paso, Texas 287
Endangered Florida Semaphore Cactus 527
Exotic pest 1267
Exotic weeds 927
Flavonoids 213

Indian Shell Rings 861
Invasive exotics 1277
lowa 605

Jalisco, Mexico 231, 637
Juarez, Mexico 287
Kansas 592

Kerala, India 809
Lakekamu basin 1071

, 473, 523, 627, 631,
637, 685, 693, LOO1, 1083, 1145
Mississippi 889, 1183
issouri 32°
a Mexico 579
orelos, Mexico 411
aout Myrsinaceae 503
Neotropics
Nevada 193, 325
New Mexico 621,
ora Mesoamericana 1145 New World 679, 7
Flora of the Venezuelan Guayana 1167 North America oe i. 325, ool, 1259
oe 377, ae 861, 1065, 1183, 1207, = North Carolina 1259,
259,12 Northern Florida 861
as ce 335 Nuevo Leén, Mexico 685
Georgia 1183 Oaxaca, Mexico 227, 473

= 9; 819,965; 937

ea

Sipa 18(4): 1309. 1999

1310

Oregon 827
Ornamental Grass 1031
ee Plants, — 579
ark Highlands 5

a Island caren Seashore 1227
Papua New Guinea 161, 751, 941,

107]
Papuasia
Peru |

1071

~

Philippines 283

Pitcher plane 1183

Pollen diameter 909

Pollen morphology of Echinopepon 479
Queretaro, Mexico 1083

Rio Nambi, Natural Reserve 389, 395, 401
Seedling development in Chamaesyce 419

Sexual ei & 1065
Sonora, Mexico 806, 1001

South Amer ao 837
South Carolina 861, 1259

1019,

Stipa 18(4)

Southeastern United States 1049, 11

Southern New Ireland 161

Southern New Mexico 987

Southern United States 1031

Tamaulipas, Mexico 800

Texas 315, 346, 350, 356, 359, 361, 365,
493,611,615, 623, 791, 800, 927,931,
1227, 1265, 1277

a/i

83, 1259

Texas Floristic Records 315, 1227
Torilis arvensis 336

United States 365, 1031, 1267
1095
Venezuelan Guayana 1167
Weed 927, 1273

Weedy 36]

West-Central Mexico, 637
Western Ghats, radia. 519, 809
Western North America 169
Western United States 647

Venezuela 401, 831,

New Names AND New COMBINATIONS

Abildgaardia papillosa Kral & M. Strong,

sp. nov. 842

Agave gracilis Garcia-Mend. & E. Martinez,

sp. nov, 227

Agave petrophila Garcia-Mend. & E.

inez nom. nov. 627
Amaranthus »

nochosp. nov. 804

Amaranthus crassipes Schltdl. var.

warnockii (I.M. Johnston) Henrickson,

87

comb. nov. 7
A 1

Henrickson, sp

nov. 8OO
oe x texensis Henrickson, nothosp.
Peaiaiann leiosolenus (Torr.) Standl. var.

sogenus (Waterf.) Spellenb. & T.

Wootten, comb. nov. 991
Anulocaulis letosolenus (Torr.) Standl. var.

howard1i Spellenb. & T. Wootten, var. nov.

989
(Lundell)

Ardisia alajuelae Pipoly &

Ricketson, comb. nov. 51
Ardisia albipedicellata Pipoly & Ricketson,

Stipa 18(4):; 1310. 1999

tucsonensis Henrickson,
4

nom. nov. 511

Ardisia eg aes ates 1) Pipoly &
Ricketson, comb. 1 15

Ardisia albise oe and a Pipoly &
Ricketson, comb. no

Ardisia azaharensis lie & Ricketson,
nom. nov. 511

Ardisia bartletit Lundell subsp. lilacina
(Lundell) Pipoly & Ricketson, comb. et
stat. nov. 466

Ardisia bristanti Pipo

jah

y & Ricketson, nom.
nov. 51

Ardisia chiriquiana ae 1) Pipoly &
Ricketsor on, comb. ne

Ardisia dressleri Pipoly & re ketson, nom.

nov. 511

Ardisia eri Pipoly & Ricketson, nom.

v.51

Aidicin nee (L . ) Pipoly &
Ricketson, comb. nov

Ardisia eciliata (Lundell) et & Ricketson,

mb. nov. 512

Ardisia ellipticifolia Pipoly & Ricketson,

nom. nov. 512

New Names AND New COMBINATIONS

Ardisia eucuneata ae Pipoly &
Ricketson, comb. nov. 512

Ardisia ae Pipoly & Ricketson, nom.
nov. 312

Ardisia Sanaa ats (Lundell) Pipoly &
Ricketson, comb. nov. 512

Ardisia gsinealenss ChundelD Pipoly &

icketson, comb. nov. 51

eater ase adel ) Pipoly &
Ricketson, comb. nov. 515

Ardisia ee ree (Lundell) Pipoly &
Ricketson,

Ardisia hornitoana Pipoly & Ricketson, nom.

comb. nov. 512

nov. 513
Ardisia hugonensis carr ) Pipoly &
Ricketson, comb. no 3
Ardisia ixcanensis (Lund ell) Pipoly &
Ricketson, comb. nov. 313
nsis (Lundell) Pipoly &
Ricketson, comb. nov. 513
Ardisia oe aaa Pipoly & Ricketson,
mb. no
Acdisia jana al ) Pipoly & Ricketson,
com
Aedista oe er Pipoly &
Ricketson, comb. n
Ardisia leptopoda angel) Pipoly &
Ricketson, comb. nc [3
Ardisia mesoamericana a Pipoly & Ricketson,

Ardisia jaliscen

4

nom. nov. 515

aes monteverdeana (Lundell) Pipoly

Ricketson, comb. 513

hae nebulosa inde Pipoly &
Ricketson, comb. n 13

Ardisia neomir ie Pipoly & Ricketson,
nom. nov. 514

Ardisia niambiensis Pipoly & Cogollo, sp.

rv. 390

Ardisia oaxacana (Lundell) Pipoly &
Ricketson, comb. nov. 514

Ardisia opegrapha Oerst. subsp. paquitensis
(Lundell) Pipoly & Ricketson, comb et stat.
nov. 457

Ardisia opegrapha Oerst. subsp. wagneri
(Mez) Pipoly & Ricketson, comb. et stat.
nov. 452

Ardisia parvidenticulata Pipoly & Ricketson,
nom. nov. 514

Sipa 18(4): 1311. 1999

Lou

Ardisia parvipunctata (Lundell) Pipoly &
Ricketson, comb. nov. 514

Ardisia samalana (Lundell) Pipoly &
Ricketson, comb. nov. 514

Ardisia steinii Pipoly & Ricketson, nom.
nov. 514

renin stevensil arg Pipoly &
Ricketson, comb. nov. 514

Ardisia toroana dande Pipoly & Ricketson,

514

Y
fol
1S)

Ardisia ea aera Pipoly &
Ricketson, comb. t

Ardisia utleyi (Lunde i) an y & Ricketson,
comb. nov. 51

Ardisia verapazensis Donn. Sm. subsp.
cucullata eee Pipoly & Ricketson,
comb. et stat. ler:

Ardisia warneri Pi or & Ricketson, nom.
nov. 515

Ardisia Itii (Lundell) Pipoly & Ricketson,

comb. nov. 515
Arnoglossum album L.C. Anderson, sp. nov.

3
Barnebydendron J.H. Kirkr., nom. nov. 817
peas aca riedelii (Tul.) J.H. Kirkr.,
comb. nov. 817
Rae pumila (Michx.) Nutt. var,
scabrella Nesom & Turner, var. nov. 499
Bulbophyllum wakoi Howcroft, sp. nov.
L019
Bulb

arajana Kral & M. Strong, sp.
nov. 4A
Bulbostylis eleocharoides Kral & M. Strong,

sp. nov. 8

Bulbostylis latifolia Kral & M. Strong, sp.

1ov. 848

— ae Kral & M. Strong,
sp. no

eee : eer Kral &
sp. nov. 852

eee: ee Kral & M. Strong,

M. Strong,

Shee : eae Kral & M. Strong,

sp. nov. 857
Clusia aristeguietae (Maguire) Pipoly, comb.

Clusia bernardoi Pipoly & Cogollo, sp. nov.
402

12
Clusta comans (Mart.)
408

Clusia deminuta Pipoly, sp. nov. 1175

Clusia divaricata (Cuatrec.) Pipoly, comb.
y

nov. 408

Clusia epiphytica (Cuatrec.) Pipoly, comb.

nov. 408
Clusia snsis (Cuatrec.) Pipoly, comb.
nov. 408

Clusia niambiensis, Pipoly, Cogollo et
396

Clusia paisarum Pipoly, s

Gonzalez, sp. nov. -

sp. nov. L179

Clusia tetragona Pipoly & Cogollo, sp. nov.
A404

Commicarpus coctoris N.A. Harriman, sp.
nov. 676

Commicarpus practermissus
Harriman, sp. nov. 681

Corsia purpurata var. wiakabui Takeuchi
& Pipoly, var. nov. 164

Crataegus ok; cane J.B.

O’Kenn rov. 178

Cratacgus a aes J.B. Phipps, sp. nov.
170

Crataegus phippsii O’Kennon, sp. nov. 185

aptec ns series Purpureofructi J.B. Phipps

N.A.

Phipps &
on, sp.

& O'Kennon, ser. nov. 184

Gatien manes indicum Hameed &
Mocinusoodat an Sp nov. 320

Croizatia | I ) Dorr, comb

Cybianthus anthuriophyllus Pipoly, sp. nov.
89

Cybiantl

s Pipoly, sp. nov. 106
Cybianthus compen anus Pipoly, sp. nov.
47
Cybianthus croatit Pipoly, sp. nov. 7 |
Cybianthus flavovirens Pipoly, sp. nov. 134
Cybianthus fosteri Pipoly, sp. nov. 144
Cybianthus grandezii Pipoly, sp. nov. 139
Cybianthus — Pipoly, sp. nov. 133
Cybiant guyanensis subsp.
Sig doe eee in Mart.) Pipoly,
comb. et stat. nov. 49
Cybianthus huampamiensis Pipoly, sp. nov
130
Cybianthus incognitus Pipoly, s 125
14 )

p. Nov.
Cybianthus jensonii Pipoly, sp. nov.

Sipa 18(4): 1312. 1999

Pipoly, comb. nov.

Sipa 18(4)

Cybianthus kayapti (Lundell) Pipoly, comb.
nov. 84

Cybianthus liesneri Pipoly & Ricketson, sp
noy. 1168

Cybianthus nestorti Pipoly, sp. nov. 59

Cybtanthus pseudolongifolius Pipoly, sp.
nov. LO1

Cybianthus timanae Pipoly, sp. nov. 53

Cybianthus vasquezii Pipoly, sp. nov. 103

Emorya rinconensis Mayfield, sp. nov. 694

Erigeron L. sect. eee anes Nesom &

oyes, sect. ~ 1183

Glochidion aie rii Takeuchi, s

Henriettea hotteana (L
comb. nov. 1026

sp. nov. 945
Urb. & Ekm.) Alain,

roe acutissima (Cuatrec.) Pipoly
icketson, comb. nov. 739
cncee callejasii (Pinel Pipoly &

icketson,

ion

comb. nov. 736

Aeneas calycosa (Hemsl.) Pipoly &

.nov. 718

Hymenandra crosbyi (Lundell) Pipoly &
Ricketson, comb. nov. 741

Hymenandra pittieri (Mez) Pipoly
Ricketson, comb. nov. 731

Hymenandra sordida oo Pipoly &
Ricketson, comb. 1

ion

Ricketson, comb

Hymenancta squamata ae Pipoly
& Ricketson, comb. 1 25

Hy ee cite lla ‘Dan Sm) Pipoly
& Ricketson, comb. 715
Hymenandra w ee (Lundell) Pipoly
Ricketson, comb. nov 728
Jasminum pipolyi Takeuchi, sp. nov. 948

Leandra humilis (Urb. & E ) Alain, comb

nov. 6
—— hybophylla (Urb.) Alain, comb.
)26
mae marigoriana Uh & Ekm.) Alain,
‘omb. . 1026
Le eae polychaeta (Urb. & Ekm.) Alain,
comb. nov. 1026
Malan alvaro Gat Ni R. Jiménez &
ichez, sp. 4
ie pastes E spejo & Lopez-Ferrart,
sp. nov. 412
Mandevilla oe ion & Mog.) J.K.
Williams, comb. 1 237

New Names AND New COMBINATIONS

oe K. Williams, sp. nov. 231
ee marck-ramuensis Takeuchi,

sp. nov. bat

Medinilla eae Takeuchi, sp. nov.
1074

SS rie a ay ‘Todsen, sp. nov. 819

Mentzelia humilus (A. Gray) Darl. var
guadalu aoe ee var. nov. 995

Muhlenbergia jaime-hintonii P.M. Peterson
& Valdés-Reyna, sp. nov. 686

Myrsine calcarata (Lundell) Ricketson &
Pipoly, comb. nov. 506

Myrsine juergensenii (Mez) Ricketson &
Pipoly, comb. nov. 506

cuibas uae Rickerson & nes sp. nov. 747

Sl tbsp. o orbiculata
(Seell) D. pales a, nov. 824
Nuphar advena (Ait.) Ait. f. subsp. ozarkana

& Stand! : D. Padgett, comb. nov.
82

cos advena (Ait.) Ait. f. subsp. ulvacea
& Standl.) D one comb. nov. 825

Nuphar pumila (Tim
oguraensis (Miki) D. a
sta. nov. 825

Nuphar pumila (Timm) DC subsp. sinensis
(Hand.-Mazz.) D. Padgett, comb. et sta
nov. 825

Nuphar sect. Astylus D. Padgett, sect. nov.

824

28

subsp.

gett, ne et

lates,

Oritrophium durangense Nesom, sp. nov.
Se ):

Padieers bolanderi var. r harfordit (Greenm.)

386

Packera buekii DK. ar & T.M. Barkley,
nom. nov. 387

2

3ar rkley. comb. nov.

Packera dimorphophylla var. intermedia
(T.M. Barkley) D.K. Trock & T.M. Barkley,
comb. nov. 386

Packera dimorphophylla var. paysoni (T.M.
Barkley) D.K. Trock & T.M. Barkley, comb.
nov. 386

ackera nec
—— D.K

nov. 386

ymexicana var.

calfei
. Trock & T.M. Baer comb.

Packera neomexicana var. toumeyi (Greene)
D.K. Trock & T.M. Barkley, comb. nov.
386

Sipa 18(4): 1313. 1999

[315

Packera pseudaurea var. flavula (Greene)
D.K. Trock & T.M. Barkley, comb. nov.
386

Packera pseudaurea var. semicordata (Mack
& Bush) D.K. Trock & T.M. Barkley, comb.
nov. 386

Packera subnuda (DC) D.K. Trock & T.M.
Barkley, comb. nov. 861

See advena Wipff & Veldkamp, sp.
nov. 1033

Phaseolus albescens McVaugh ex R. Ramirez
A Delgado, sp. nov. 638

Physaria iene A. Gray var. yesicola
Sivinski, var. nov. 673

Pilea hedemarkii Takeuchi, sp. nov. 770

Pinus greggii Engelm. ex Parl. var. australis
Donahue & Lopez, var. nov. 1092

Polianthes oe lano & Garcia-Mend
p 73

Mend.,
ee ate Henrickson & T.Van
evender, sp. nov. LOO2
Prunus gideonii Takeuchi, sp. nov. 765
Psychotria howcroftii Takeuchi, sp. nov.
Psychotria matambuai Takeuchi, sp. nov.
a0

Psychotria osiana Takeuchi & Pipoly, sp.
nov. 162

Ruellia jimulcensis Villarreal, sp. nov. 223

Sagraea abbottii (Urb.) Alain, comb. nov.
1027

Sagraea barahonensis (Urb. & Ekm.) Alain,
comb. nov. 1027

Sagraea cinerea (Cogn.) Alain, comb. nov.
2

ila(Urb. & Ekm.) Alain, comb

“SO r
nov. 1027
Sagraea ellipsoidea (Urb. & Ekm.) Alain,
comb. nov. 1027

Sagraea fuertesii (Cogn., Urb.) Alain, comb.
nov. 1028

Sagraea gracilis (Alain) Alain, comb. nov.
1028

Sagraea lanceifolia (Urb.) Alain, comb. nov.
1028

Sagraea oligantha (Urb.) Alain, comb. nov.
1028

1314

Sagraea polychaete (Urb. & Ekm.) Alain,

comb. nov. L028

Sagraea pusilliflora (Cogn., A. & C.DC.)
Alain, comb. nov. 1028

Sagraea setulosa (Urb.) Alain, com
1028

ion

nov.

] ii (J ld & Skean) Alain, comb

Sarracenia rosea f. luteola (Hanrahan &
Miller) Naczi, EW. Case, & R.B. Case, comb.
nov. 1192

Sarracenia rosea Naczi, EF. W. Case, & R.B.
Case, sp. nov. 1188

Setaria subgenus Reverchoniae W.E. Fox

59

subgenus nov. 103

Setaria seyecchonil (Vasey) Pilg. subsp.
firmula (Hitche. & Chase) W.E. Fox, comb.
nov. 1042

Setaria reverchonii (Vasey) Pilg. subsp

ramiseta (Scribn.) W.E. Fox, comb. nov.

1041

Setaria utowanaea (Scribn.) Pilg. var.
Fox, comb. nov.

ophiticola (Léon) W.E.
1043

Sipa 18(4); 1314. 1999

Stipa 18(4)

Setaria utowanaea (Scribn.) Pilg. var.
subtransiens (Hitchc. & Ekman) W.E. Fox,
comb. 1043

Spiraea « hitchocockit W. Hess & N.
Stoynoff, hybrida nov. 827

Sporobolus gaan Weakley & P.M.
Peterson, sp. nov. 258

Stylogyne viridis ag Ricketson &
Pipoly, comb. nov. 11

Trachypogon mayaénsis es &S.D. Jones,
sp. nov. 242

Trichospermum stevensii Takeuchi, sp. nov.

y77

Tripogon sivarajanii Sunil, sp. nov. 809

Tripogon vellarianus Pradeep, sp. nov. 811
piu eae Clark, gen. nov. 632

rcuttii (Greenm.) B.L. Clark,
comb. nov. 633

Xanthostemon fruticosus Peter G.

son & L. Co

Wil-

), sp. nov. 283

SIDA GUIDELINES FOR CONTRIBUTORS

SIDA, CONTRIBUTIONS TO BOTANY is an international journal of systematic botany contain-
ing primary research papers sensu lato, including anatomy, ec iemae chemotaxonomy,

cladistics, ecology, floristics, genetics & evolution, numerical taxonomy, paleobotany, and
palynology. Si1pa is open to all don anywhere; coverage is not Anes to any geo-
graphical area.

All manuscripts submitted to Sip are considered by at least two reviewers. Manuscripts
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817-33 12 fax. Our electronic mail address is sida@brit.org. Access our SIDA home
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Manuscripts must be double spaced throughout and three copi nt to the editor. Upon

acceptance for publication, please return two copies along si a copy in your word-pro-
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double hyphens (6--8). Foot notes, figure legends, appendices, and tables should be on
separate pages and inserted at end of manuscript following references.

Abstract

Every paper should include both an English and Spanish, or another major language ab-
stract

Specimen Citations

Use the collector’s last name and number or (s.n.) and underline. Abbreviate the months of

the year by the first three letters of the month without a period. Example: Texas. Jeff Davis
Co.: | mi W of Fort Davis, 3 Jan 1972, Smith 118 (BRIT)

Author Citations

Use the author abbreviations as given in the Author A bbreviati npiled at the Herbarium,
Royal Botani ardens, Kew (1992). Use et or the ampersand (&) between two author
for more ae two authors, restrict to the first one followed by et al. Do not ees or
underline in the manuscript the terms et, et al., ex, or in

Latin Descriptions or Diagnoses; English Descriptions

Please provide a Latin diagnosis for each new taxon. After the Latin diagnosis, an English
or Spanish version is recommended highlighting the diagnostic characters. Write plant
descriptions as incomplete sentences with phrases separated by semicolons.

Figures

Please present ALL figures as a unified, single sequence of figures. A scale should be in or
on the illustration or photograph; reduction will automatically reduce the scale and sub-
ject proportionately. In halftones, sharp glossy photographs with good contrast are neces-

Sipa 18(4): 1315. 1999

1316 Sipa 18(4)

sary for good reproduction. Please mount illustrative material (halftones, line drawings,
etc.) on flexible paper so that may be mounted and on a high-speed drum scanner. You
may also submit graphic materials in electronic format. Please consult the editor for proper
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References

This includes all of the literature cited in the text and may include other article citations
the author may deem desirable. Normal text references should be cited as follows: ‘Ricketson
and Pipoly (1997) stated...’ or ‘the latest revision (Ricketson & Pipoly 1997) when refer-
ence is used as authority for a statement. When there are three or more awe use only
the name of the first author followed by et al.: ‘Barrie et al. (1992a) stated’. References at
the end of the article are arranged alphabetically and chronologically ses use of a,b,
etc. if an author had more than one publication ina given year. Author's names are typed

in lower case except for initial capitals. Only the firse letter of the initial word, proper

nouns, and proper adjectives of titles should be aie For journal eee use
Botanico-Periodicum-Huntianum. Reterence examples:
a oe

Barrie, ER., CLE. ee anc . Reveal. 1992a. The need to change Article 8.3 of the

one Taxon 41:508-

Ricketson, J. a ae aoe 1997. A synopsis of the genus Gent/ea (Myrsinaceae) and a key
to the genera of Myrsinaceae in Mesoamerica. Sida 17:697--707.

Crawford, D.J. 1983. Phylogenetic and systematic inferences from electrophoretic stud-
ies. In: S.D. Tanksley and T.J. Orton, eds. Isozymes in plant genetics and breeding, Part
A. Elsevier, Amsterdam. Pp. 257--28

Sivinski, R.C. and K. aes (eds.). 1995. Inventory of rare and endangered plants of

New Mexico. New xico Forestry and Resources Conservation Division, ns Min-
erals and Natural Sone Department. Misc. Pub. No. 4.
Abbreviations

When the following abbreviations are used the period ts See Distance: mm, dm, cm,
, km, ft, mi; directions (in caps): S, N, E, W, months: first 3 letters only, Jan, Feb, ete.
ae ple: Potter Co.: 2 km W of one 5S Jun ee , Smith L18 (SMU)

Documented Plant Chromosome Numbers

Refer to Sipa 18(3):909-925, 1999,

Sipa 18(4): 1316. 1999

CURRENT RESEARCH IN

ba

ayy

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privately by Lloyd
H. Shinners, above
left in 1962. It was
inherited by
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Director Emeritus of
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multiple-topic

format.

ss

Sida, Contributions to Botany (SCB) has been a source of currer
research in classical and modern systematic botany for reade
throughout the world for nearly 40 years. It contains primar
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obotany, palynology, and phylogenetic systematics. Coverac

is global: it is not restricted to any geographicel area, an
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dished

Each issue contains articles on various groups of plants, suc

as palms, ferns, lilies, irises, legumes, primroses morning-glorie

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appear in English or Spanish, with abstracts in both language
All papers are peer reviewed and are frequently illustrate
with maps and line drawings, frequently with full color. Eac

issue also includes short communications on floristic disco
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A new species of Phaseolus (Fabaceae) from West-Central Mexico
Taxonomy of Cyphomeris (Nyctaginaceae) based on multivariate
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results from the 1995 Bismarck-Ramu Expedition in Papua New Guin
Studies in New World Amaranthus (Amaranthaceae)

Eight novelties in Abildgaardia and Bulbostylis (Cyperaceae)
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Two new species of Tripogon (Poaceae) from India

Ethnobotany serving society: A case study from the
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3M has been published since 1987 by BRIT as an adjunct to SCB,
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a poth

ma
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Be
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a
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a
aw
wr

st

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Index to new names and new combinations in this issue

Anulocaulis leiosolenus (Torr.) Standl. var. gypsogenus (Waterf.) Spellenb. & ‘T. Wootten, comb. nov. 992
Anulocaulis leiosolenus (Torr.) Standl. var. howardii Spellenb. & T. Wootten, var. nov. 989
Ardisia verapazensis Donn. Sm. subsp. cucullata (Lundell) Pipoly & Ricketson, comb. et stat. nov. 1156

Bulbophyllum wakoi Howeroft, sp. nov. 1019

Clusia deminuta Pipoly, sp. nov. 1175
Clusia paisarum Pipoly, sp. nov. 1179
Cybianthus liesneri Pipoly & Ricketson, sp. nov. 1168

Erigeron |.. sect. Geronpternix Nesom & Noyes, sect. nov. 1163
Glochidion beehlerii Takeuchi, sp. nov. 945
Henriettea hotteana (Urb. & Ekm.) Alain, comb. nov. 1026

Jasminum pipolyi Takeuchi, sp. nov. 948

Leandra humilis (Urb. & E-km.) Alain, comb. nov. 1026
Leandra hybophylla (Urb.) Alain, comb. nov. 1026
Leandra marigotiana (Urb. & Ekm.) Alain, comb. nov. 1026
Leandra polychaeta (Urb. & Ekm.) Alain, comb. noy. 1026

Medinilla bismarck-ramuensis Takeuchi, sp. nov. 1071
Medinilla sapoi-riverensis Takeuchi, sp. nov. L074
Mentzelia humilus (A. Gray) Darl. var guadalupensis Spellenb., var. nov. 995
Pennisetum advena Wiptf & Veldkamp, sp. nov. 1033

inus greggii Hngelm. ex Parl. var. australis Donahue & Lopez, var. nov. 1092
Portulaca yecorensis Henrickson & T.Van Devender, sp. nov. 1002
Psychotria matambuai Takeuchi, sp. noy.

950
Sagraea abbottii (Urb.) Alain, comb: nov. 1027
Sagraea barahonensis (Urb. & [-km.) Alain, comb. nov. 1027

Sagraea cinerea as a cou nov. 1027

Sagraea curvipila (Urb. & E:km.) Alain, comb. nov. 102
Sagraea ellipsoidea (Urb. & Bien) Alain, comb. nov. 1027
Sagraea fuertesii (Cogn., Urb.) Alain, comb. nov. 1028
Sagraea gracilis (\lain) Alain, ‘comb. nov. 1028

— ane (Urb.) Alain, comb. nov. 1028

Sagra gantha (Urb.) Alain, comb. nov. 1028

nae pee (Urb. & Ekm.) Alain, comb. nov. 1028
Sagraea pu ora (Cogn., A. & C.DC.) Alain, comb. nov. 1028
Sagra see Irb.) Alain, comb. nov. 1028

Sagraea er ae & Skean) Alain, comb. nov. 1029

Sarracenia rosea Naczi, I'.\W. Case, & R.B. Case, sp. nov. 1188

Sarracenia rosea f. intel (Hanrahan & Miller) Naczi, I .W. Case, & R.B. Case, comb. nov. 1192
Setaria subgenus Reverchoniae WE. Fox, subgenus nov. 1039

Setaria reverchonii (Vasey) Pi - subsp. firmula (Hitche. & Chase) W.E. Fox, comb. nov. 1041
Setaria reverchonii (Vascy) Pilg. subsp. ramiseta (Scribn.) W.L. Fox, comb. nov. 1041

Setaria utowanaea (Scribn.) Pilg. var. ophiticola (Léon) W.E. Fox, comb. nov. 1043

Setaria utowanaea Gatiba) Pilg, var. subtransiens (Hitchc. & Ekman) W.E. Fox, comb. 1043

Stylogyne viridis (Lundell) Ricketson & Pipoly, comb. nov. 117
) I

Trichospermum stevensii Takeuchi, sp. nov. 1077

Novelties in the Myrsinaceae from the Venezuelan Guayana

Jobn J. Pipoly Ul and Jon M. Ricketson

1167

New species of Clusia (Clusiaceae) from the Cordillera Occidental of Colombia
Jobn Jf. Pipoly Mand Alvaro Cogollo P

1175

Sarracenia rosea (Sarraceniaceac), a new species of pitcher plant from the southeastern United States
Robert F.C. Naczi, Eric M. Soper, Frederick W. Case, Jr, and Roberta B. Case

1183

The vascular flora of Biscayne National Park, Vlorida

Richard Stalter, Josef Tamory, Patrick Lynch, and Brian Lockwood

1207

A checklist for the vascular plants of Padre Island National Seashore
LG. Negrete, A.D. Nelson, |. R. Goetze, L. Macke, T. Wilbtan, and A, Day
1297

An annotated preliminary checklist of the dicotyledonous lianas and vines trom the Las Cruces Biological Station,
Costa Rica
Alexander Krings

947

Gamochaeta simplicicaulis (Asteraceae: Gnaphalicae) in four southeastern states and new for North Americ:
Guy L. Nesom
1259
Vicia lutea (l'abaceae) new to Texas
Amanda K. Neill
265
cia ee (C a ie), another oo tal exotic pest in the United St:
Kathleen daniel F. Austi

267
Paederia foetida (Rubiaceac), new to the Hora of North Carolina
Pete Diamond

Lapsana communis (Asteraceae) new for Texas and notes on invasive exotics
Ruben O’Kennon, Theodore M. Barkley, George M. Diggs, Jr. and Barney Lipscomb
1277

Bouteloua rigidiseta (Poaceae) new to Louisiana
Charles M. Allen, Chris 8. Reid, and Chris H. Doffitt
285
Book notices and reviews L000, 1008, 1024, 1030, 1048, 1064, 1082, 1094, 1166, 1246, 1284, 1286
Reviewers for volume 18, 1998-1999 1295

Index to volume 18, 1998-1999 1296

mn

Sida guidelines for contributors [3

an?

Sida Reference Guide 131

ei > | (continued on inside back cover)

4 GE ISSN 0036-1488